1.

5 WATER MOVEMENT MAY BE GOVERNED BY DIFFUSION OR BY

BULK FLOW
One objective of plant physiology is to understand the dynamics of water as it flows into and out
of cells or from the soil, through the plant, into the atmosphere. Movement of substances from
one region to another is commonly referred to as translocation.Mechanisms for translocation may
be classified as either active or passive, depending on whether metabolic energy is expended in
the process. It is sometimes difficult to distinguish between active and passive transport, but the
translocation of water is clearly a passive process. Although in the past many scientists argued for
an active component, the evidence indicates that water movement in plants may be indirectly
dependent upon on expenditure of metabolic energy. Passive movement of most substances can
be accounted for by one of two physical processes: either bulk flow or diffusion. In the case of
water, a special case of diffusion known as osmosis must also be taken into account.
1.5.1 BULK FLOW IS DRIVEN BY HYDROSTATIC PRESSURE
Movement of materials by bulk flow (or mass flow) is pressure-driven. Bulk flow occurs when an
external force, such as gravity or pressure, is applied. As a result, all of the molecules of the
substance move in a mass. Movement of water by bulk flow is a part of our everyday experience.
Water in a stream flows in response to thehydrostatic pressure established by gravity. It flows
from
the faucet in the home or workplace because of pressure generated by gravity acting on standing
columns of water in the municipal water tower. Bulk flow also accounts for some water movement
in plants, such as through the conducting cells of xylem tissue or the movement of water into
roots. In Chapter 9, we discuss how bulk flow is a major component of the most widely accepted
hypothesis for transport of solutes through the vascular tissue.
1.5.2 FICK’S FIRST LAW DESCRIBES THE PROCESS OF DIFFUSION
Like bulk flow, diffusion is also a part of our everyday experience. When a small amount of sugar is
placed in a cup of hot drink, the sweetness soon becomes dispersed throughout the cup. The
scent of perfume from a bottle opened in the corner of a room will soon become uniformly
distributed throughout the air. If the drink is not stirred and there are no mass movements of air
in the room, the distribution of these substances occurs by diffusion. Diffusion can be interpreted
as a directed movement from a region of a high concentration to a region of lower concentration,
but it is accomplished through the random thermal motion of individual molecules (Figure 1.4).
Thus, while bulk flow is pressure-driven, diffusion is driven principally by concentration differences.
Diffusion is a significant factor in the uptake and distribution of water, gases, and solutes
throughout the plant. In particular, diffusion is an important factor in the supply of carbon dioxide
for photosynthesis
as well as the loss of water vapor from leaves. The process of diffusion was first examined
quantitatively by A. Fick. Fick’s first law, formulated in 1855, forms the basis for the modern-day
quantitative description of the process.
J = − D · A · _C · l− 1 (1.1)
J is the flux or the amount of material crossing a unit area per unit time (for example, mol m− 2 s−
1). D is the diffusion coefficient, a proportionality constant that is a function of the diffusing
molecule and the medium through which it travels. A and l are the cross-sectional area and the
length of the diffusion path, respectively. The term _C represents the difference in concentration
between the two regions, also known as the concentration gradient. _C is the driving force for
simple diffusion. In the particular case of gaseous diffusion, it is more convenient to use the
difference in density (gm m− 3) or vapor pressure (KPa, kilopascal) in place of concentration. The
negative sign in Fick’s law accounts
for the fact that diffusion is toward the lower concentration or vapor pressure. In summary, Fick’s
law tells us that the rate of diffusion is directly proportional to the cross-sectional area of the
diffusion path and to the concentration or vapor pressure gradient, and it is inversely proportional
to the length of the diffusion path.