ONE

Water & It's Role in Plants

This chapter deals with the ecological and physiological importance of water, its various roles in
plants, Its peculiar properties, how it occurs in cells and tissues, and the forces Involved In Its
movement. The terminology of cell water relations also will be discussed.

The Importance of water

The importance of water will be treated in terms of its ecological and physiological roles. Much
additional material can be found in the first three volumes of Ruhland's "Encyclopedia of Plant
Physiology," also In Crafts et al. (1949), Hagan et al. (1967), Kozlowski (1964), Rutter and
Whitehead (1963), Slatyer (1967), and Slavik (1965).

Ecological importance of water

The distribution of vegetation over the surface of the earth is controlled more by the availability
of water than by any other single factor. Regions where rainfall is abundant and falrly evenly
distributed over the growing season have lush vegetation. Examples are the rain forests of the
tropics, the vegetation of the Olympic Peninsula of the Northwestern United States, and the
luxuriant cove forests of the Southern Appalachians. Where summer droughts are frequent and
severe, forests are replaced by grasslands, as in the Steppes of Asia and the prairies of North
America. Further decrease in rainfall results in semidesert with scattered shrubs, and finally
deserts.

Even the effects of temperature are partly exerted through water relations because decreasing
temperature is accompanied by decreasing rates of evaporation and transpiration, and
increasing temperature is accompanied by increasing rates. Thus, an amount of rainfall
adequate only for grasslands
HEREDITARY POTENTIALITIES ENVIRONMENTAL FACTORS

Depth and extent of root systems SOIL Texture, structure, depth, chemical

Size, shape and total area of composition and pH, aeration, temperature,
leaves, and ratio of internal waterholding capacity, and water conductivity
to external surface

Number, location, and behavior of ATMOSPHERIC. Amount and distribution of

Stomata precipitation.
Ratio of precipitation to evaporation
Radiant energy, wind, vapor pressure and
other factors affecting evaporation
and transpiration

PLANT PROCESSES AND CONDITIONS

Water absorption
Ascent of sap
Transpiration
Internal water balance as reflected in water potential, turgidity, stomatal
opening, and cell enlargement.
Effects on photosynthesis, carbohydrate and nitrogen inetabolism, and other
metabolic processes

QUANTITY AND QUALITY OF GROWTH

Size of cells, organs, and plants

Dry weight, succulence, kinds and amounts of various compounds

produced and accumulated

Root/shoot ratio

Vegetative versus reproductive growth

FIG. 1.1 Diagram showing how the quantity and quality of plant growth is controlled by
hereditary and environmental factors operating through the internal processes and conditions
of the plant, with special reference to factors affecting water relations.

in a hot climate can support forests in a cooler climate where the rate of evapotranspiration is much
lower. This fact was responsible for development of the concept of the rainfall-evaporation ratio by
Transeau (1905) and for the Inclusion of data for potential evapotranspiration along with temperature in
formulas from which life zones are predicted from climatic data (Holdridge, 1962). Furthermore,
temperature may affect life processes as much by its effects on the properties of water as by its effects
on rates of chemical reactions (Tanford, 1963).

Physiological importance of water

The ecological importance of water is the result of its physiological importance. The only way in
which an environmental factor such as water can affect plant growth is by affecting Internal
physiological processes and conditions. The interrelationships between plants and their
environment with respect to water are summarized in Fig. 1.1.

Almost every plant process is affected directly or indirectly by the water supply. Many of these
effects will be discussed later, but it can be emphasized here that within limits metabolic activity
of cells and plants is closely related to their water content. For example, the respiration of
young, maturing seeds is quite high, but it decreases steadily during maturation as water
content decreases (see Fig. 1.2). Incidentally, the decrease in water content during maturation
of seeds encased in fleshy fruits, such as tomato, is difficult to explain (Mcilrath et al., 1963).
The respiration rate of air-dry 'seeds is very low and increases slowly with increasing water
content up to a critical point, where there is a rapid increase in respiration with further increase
in water content (Fig. 1.3). The practical importance of the relationship between water content
and respiration is shown in connection with storage of grain, because high water contents result
in "heating" and spoilage.

Growth of plants is controlled by rates of cell division and enlargement and by the supply of
organic and inorganic compounds required for the synthesis of new protoplasm -and cell walls.
Cell enlargement is particularly dependent on at least a minimum degree of cell turgor, and
stem and leaf elongation are quickly checked or stopped by water deficits (Loomis, 1934; Miller,
1965; Thut and Loomis, 1944).
Decrease in water content Invariably reduces the rate of photosynthesis (Brix, 1962) and usually
reduces the rate of respiration. However, with decreasing water content, drying. pine needles
and a few other kinds of tissue sometimes show a temporary increase in respiration before the
final decrease and death (Brix, 1962; Montfort and Hohn, 1950; Parker, 1952). In summary,
decreasing water content is accompanied by loss of turgor and wilting, cessation of cell
enlargement, closure of stomata, reduction in photosynthesis, and interference with many basic
metabolic processes. Eventually, continued dehydration causes disorganization of the
protoplasm and death of most organisms. The effects of water deficits on physiological
processes are dis cussed in more detall In Chap. 10.

Uses of water in plants

The importance of water can be summarized by mentioning its most important functions, which
will be grouped under four general headings.

A CONSTITUENT OF PROTOPLASM. Water is as Important quantitatively as qualitatively,

constituting 80 to 90 percent of the fresh weight of most herbaceous plant parts and over 50
percent of the fresh weight of woody plants. Some data on water content of various plant
structures are shown in

Table 1.1. Water is just as important a part of the protoplasm as the protein molecules which
constitute the protoplasmic framework, and reduction of water content below some level 's
accompanied by changes in structure and ultimately in death. It is true that a few plants and
plant organs can be denydrated to the air-dry condition, or even to the oven-dry condition in
the case of some kinds of seeds and spores, without loss of viability, but a marked decrease in
physiological activity always accompanies decrease in water content.

SOLVENT. A second essential function of water in plants is as the solvent In which gases,
minerals, and other solutes enter plant cells and move from cell to cell and organ to organ. The
permeability of most cell walls and membranes to water results in a continuous liquid phase
extending through out the plant in which translocation of solutes of all kinds occurs.

REAGENT. Water is a reactant or reagent in many important processes, including photosynthesis

and hydrolytic processes such as the hydrolysis of starch to sugar. It is just as essential in this
role as carbon dioxide or nitrate.
MAINTENANCE OF TURGIDITY. Another essential role of water is in the maintenance of the
turgidity so essential for cell enlargement and growth, and for maintaining the form of
herbaceous plants. Turgor also is important in the opening of stomata and the movements of
leaves, flower petals, and various specialized plant structures. Inadequate water to maintain
turgor results in an immediate reduction of vegetative growth.

Properties of water

The Importance of water in living organisms results from its unique physical and chemical
properties. Bernal (1965) comments that nearly all of its properties are anomalous. The
importance of these properties in life processes has been recognized for several decades and
discussed by many writers from Henderson (1913), Bayliss (1924), and Gortner (1938) to
Hutchinson (1957, pp. 195-200). Important collections of papers were published in Symposium
no. 19 of the Society for Experimental Biology and in vol. 125, art. 2 of the Annals of the New
York Academy of Sciences. The properties of water and solutions also are discussed in some
detail in chap. 1 of Siatyer. (1967).

Water plays such a special role in living organisms that we can scarcely imagine life
without it In fact, the absence of water on the moon and planets would be regarded as
sufficient evidence that life as we know it cannot exist there.

Isotopes of water

The three isotopes of hydrogen having atomic weights of 1, 2, and 3 make it possible to
differentiate "racer water" from ordinary water. In the 1930s “heavy water," water containing
deuterium (hydrogen of atomic weight 2), became available and was used widely in biochemical
studies. It was also used extensively in studies of permeability of animal and plant membranes
(Ordin and Kramer, 1956; Ussing, 1953, for example). However, in recent years deuterium has
been largely supplanted as a tracer by water containing tritium (hydrogen of atomic welght 3),
for example, in the experiments of Raney and Vaadla (1965). Tritium is radioactive and therefore
more convenlent as a label, being easier to detect than deuterium, which requires use of a mass
spectrometer.

A stable isotope of oxygen with an atomic weight of 18 makes it possible to study the role of the
oxygen in water. An example is the series of experiments with H,180 which demonstrated that
the oxygen released during photosynthesis comes from water rather than from carbon dioxide,
as was previously supposed (Ruben et al., 1941).
Unique properties of water

Data on the chemical and physical properties of water were collected recently by Kavanau
(1964) and Kohn (1965). A few of its more unique properties will be discussed. Water has the
highest specific heat of any substance except liquid ammonia, which is about 13 percent higher.
The standard unit for measuring heat, the calorie, is the amount of energy required to warm
one gram of water one degree. The heat of vaporization is the highest known, 540 cal/g at
100°C, and the heat of melting, 80 cal/g, Is also unusually high. The high specific heat of water
tends to stabilize temperatures and is reflected in the uniform temperature of Islands and land
near large bodies of water. The high heat of vaporization means that evaporation of water has
pronounced cooling effect and condensation has a warming effect. Water likewise is an
extremely good conductor of heat compared with other liquids and nonmetallic solids, although
it is poor compared with metals.

Water is falrly transparent to visible radiation. It allows light to penetrate and makes it
possible for algae to carry on photosynthesis and grow to considerable depths in bodies of
water. It is opaque to long wavelengths, so that water filters are fairly good heat absorbers (see
Fig. 1.4).

Water has a much higher surface tension and viscosity than most other liquids because
of the high Internal cohesive forces between molecules. This also provides the tensile strength
required by the cohesion theory of the ascent of sap. It has a density which is exceeded among
liquids only by

FIG. 1.4 Transmission of radiation of varians wavelengths through layers of water of different
thicknesses. The numbers on the curves refer to the thickness of the layers. Note that
transmission is hich in shorter wavelengths and decreases for longer wavelengths. (After
Hollaender, 1956, vol. 3, p. 195.)

molten metals such as mercury and is remarkable in having its maximum density at 4°C instead
of at freezing. Even more remarkable is the fact that water expands on freezing, so that ice has a
volume about 10 percent greater than the liquid water from which it was formed (see Fig. 1.5).
This explains the fact that ice floats and that pipes and radiators burst when the water in them
freezes. Incidentally, if ice sank, bodies of water in the cooler parts of the world would all be
filled permanently with ice.

Water is very slightly ionized; only one molecule in 5.5 x 108 is dissociated. Since a hydrogen lon
is a bare proton and cannot exist alone, it becomes associated with a water molecule to form a
hydronlum lon. The dissociation of water can therefore be described as
2H2O→H3O+ + OH-.

Because it is so little lonized, water has a high dielectric constant which contributes to its
behavior as an almost perfect solvent. It is a good solvent of electrolytes because the attraction
of ions to the positive and negative charges on water molecules forms dipole bonds. As a result,
each ion is surrounded by a shell of water molecules which keeps ions of opposite

charge separated. It is a good solvent for most nonelectrolytes because it forms hydrogen bonds
with amino and carboxyl groups. It tends to be adsorbed or bound strongly to the surfaces of
cellulose, clay micelles, protein molecules, and many other substances. This characteristic is of
great importance in soil and plant water relations.

Explanntion of unique properties of water

The unusual combination of properties which makes water so important ecologically and
physiologically has been recognized for several decades (Bayliss, 1924; Gortner, 1938;
Henderson, 1913). Although the idea was proposed by Latimer and Rodebush (1920), this
unique combination of properties has only recently been explained by assuming that water
molecules are assoclated in an orderly structure by hydrogen bonding (Bernal, 1965: Crafts et
al., 1949; Kavanau, 1964; Pauling, 1960; Tanford, 1963).

To make this explanation clearer we shall review briefly the kinds of forces which
operate among atoms and molecules. First are the ordinary chemical bunds such as the covalent
bonds between oxygen and hydrogen in water or carbon and hydrogen in methane. These are
broken during chemical reactions, as when starch is hydrolyzed to sugar. If covalent bonds were
the only typo, there would be no liquids or solids, because covalent bonding does not hold
molecules together. However, there are various physical or electrical forces, such as the effects
of dipoles, van der Waals' forces, and hydrogen bonds, which operate between adjacent
molecules. Some molecules are polar' or electrically unsymmetrical because they have
positively and negatively charged areas which attract one molecule to another. Water molecules
with a structure show this dipole effect rather strongly, while substances such as methane or
carbon tetrachloride do not because they have no unsymmetrical distribution of electrical
charges.

There is a tendency of the positively charged nucleus of a moleculo to attract the negatively
charged electrons of neighboring molecules. These attractions, called van der Waals' forces, are
rather weak and are effective only if the molecules are close together. Liquefaction of gases
depends on van der Waals forces holding the molecules together, and the bolling point of most
liquids therefore depends on the energy required to break the van der Waals' forces. Liquefied
gases evaporate rapidly because Physical properties such as boiling point, heat of vaporization,
viscosity, are so weakly bound together. vaporate rapidly because the molecules and surface
tension depend on the strength of Intermolecular bonding. viously, the peculiar properties of
water depend on intermolecular forces much stronger than those already mentioned. These
attractive forces are supplied by hydrogen bonds, which result from attraction of the hydrogen
atoms of one water molecule to the oxygen atoms of adjacent molecules. The forces thus
produced by the peculiar distribution of charges on water molecules (Bernal, 1965; Tanford,
1963) bind water molecules in the sym metrical lattice structure shown diagrammatically in Fig.
1.8. In Ice, all water molecules are arranged in a lattice with unusually wide spacing. This
arrangement results in a density less than that of liquid water.

As Ice melts, about 15 percent of the bonds break, and perhaps 8 percent of the molecules
escape from the lattice. This results in a partial collapse of the lattice and an increase to
maximum density at 4°C. As the temperature rises above 4º, increase in thermal agitation of the
molecules produces an increase in volume (see Fig. 1.5). There is some uncertainty about the
exact structure of liquid water, and some Investigators propose that It consists of flickering
clusters" or "icebergs" of structured molecules separated by regions of randomly arranged
molecules (Kavanau, 1964). In any event, It seems that the structure is only partly destroyed by
Increase in temperature and at least 70 percent of the hydrogen bond energy found in Ice
remains at 100°C.

The structure is somewhat modified by the pH, which affects the distance of the hydrogen from
the oxygen atoms, and by lons, because of their attraction for water molecules, lons form dipole
bonds with water molecules. The result is that the ions become surrounded by shells of water
molecules which are firmly bound (Fig. 1.7). In fact, Bernal (1965) describes ions, protein

FIG. 1.7 Arrangement of water molecules in oriented layers or shells around ions with the
hydrogen fons nearest to cations, the oxygen ions nearest to anions. These she is rend to
separate ions of opposite charge and enable them to exist in solution. They also disrupt the
normal structure of water and slightly increase the volume. (After Buswell and molecules, and
cell surfaces in general as being coated with "ice," l.e., with layers of structured water
molecules, Solution of alcohols, amides, and other polar organic liquids in water results in a
more strongly structured system than occurs in the separate substances. This is shown in the
high viscosity of such solutions. For example, the viscosity of an alcohol-water mixture at 0°C
tour times that of water or alcohol alone. However, this structure is easily broken by high
temperatures (see Fig. 1.8). The addition of nonpolar substances such as benzene or other
hydrocarbons to water breaks hydrogen

us and produces "holes" or disorganized areas in the structure which are ounded by areas with
a tighter structure. The water bound on large molecules such as proteins has an important
effect on their structure; and Tanford (1963) cites evidence suggesting that the relative stability
of the structure of viruses, DNA, and globular proteins is determined by the stability of the
water associated with them.

It was mentioned earlier that the changes in volume of water during freezing and
thawing are caused by changes in the proportion of water molecules bound in an organized
lattice by hydrogen bonds. The high boiling point results from the large amount of energy
required to break hydrogen bonds, and two must be broken for each molecule evaporated.
Methane (CH) has nearly the same molecular weight as water, but it boils at -161°C because no
hydrogen bonds occur and only a small amount of energy is required to break the van der
Waals' forces holding the molecules together in the liquid.

The unusually high viscosity and surface tension also result from the fact that hydrogen bonds
between water molecules resist rearrangement. Water wets and adheres to glass, clay micelies,
cellulose, and other substances which have exposed oxygen atoms at the surface with which
hydrogen bonds can be formed. It does not wet paraffin and other hydrocarbons because it
cannot form hydrogen bonds with them. Water wets cotton, because it forms numerous
hydrogen bonds with oxygen atoms of the cellulose molecules, but not nylon, which has few
atoms with which hydrogen bonds can be formed.

It has been suggested by Drost-Hansen (1965) and others that there are discontinuitles In the
properties of water which are of biological Importance. Drost-Hansen claims that there are
anomalies in the properties of water at 15, 30, 45, and 60°C which are reflected in the behavior
of living organisms. This clalm has been received rather sceptically and Falk and Kell (1966)
attribute the alleged discontinuities to errors to measurements.

Properties of aqueous solutions

In plant physiology we seldom deal with pure water because the water in plants and in their
root environment contains a wide range of solutes. Therefore, it is necessary to understand how
the properties of water in solution differ from those of pure water. Only a brief discussion is
possible here, and readers are referred to physical chemistry texts for a full development of
these Ideas. The characteristics of water in solution can be shown concisely by tabulation of its
colllgative properties, i.e. the properties associated with the concentration of solutes dissolved
in it. These are shown in Table 1.2 and include the effects on vapor pressure, boiling and
freezing points, and osmotic pressure.

VAPOR PRESSURE. The decrease in vapor pressure of water in solution essentially the result of
its dilution by the addition of solutes. This is Snown by Raoult's law, which states that the vapor
pressure of solvent vapor in equilibrium with a dilute solution is proportional to the mole
fraction of solvent in the solution: (1.1)

where e is the vapor pressure of the solution, eº the vapor pressure of pure solvent, the
number of moles of solvent, and n, the number of moles of