Movement of water and solutes – Chapter 30
 Transport in plants occurs on three levels:
(1) the uptake and loss of water and solutes by individual cells
(2) short-distance transport
of substances from cell to
cell at the level of tissues
or organs
(3) long-distance transport
of sap within xylem and
phloem at the level of
the whole plant
***We will focus on this level
The root-soil interface
 Water moves through soil by bulk flow (hydrostatic pressure)
 At root interface, movement changes to diffusion
 Water and solutes enter most actively near root tip, root hairs enhance uptake
Water transport
 Three routes are available for lateral transport, the movement of water and
solutes from one location to another within plant tissues and organs
 Transcellular
 Symplast
 Apoplast
Water movement
 In one route, substances move out of one cell, across the cell wall, and into
the neighboring cell, which may then pass the substances along to the next
cell by same mechanism
 This transmembrane route requires repeated crossings of plasma
membranes
 The second route, via the symplast, requires only one crossing of a plasma
membrane
 After entering one cell, solutes and water move from cell to cell via
plasmodesmata
 The third route is along the apoplast, the extracellular pathway consisting of
cell wall and extracellular spaces.
 Water and solutes can move from one location to another within a root or
other organ through the continuum of cell walls before ever entering a cell
Root pressure
 Ions actively pumped into xylem de
xylem
 Increased pressure forces fluid up stem; very limited in distance (50 cm max)
Introduction
 Xylem sap flows upward to veins that branch throughout each leaf, providing
each with water
 Plants lose an astonishing amount of water by transpiration, the loss of water
vapor from leaves and other aerial parts of the plant
 An average-sized maple tree losses more than 200 L of water per hour
during the summer
The ascent of xylem sap depends mainly on transpiration and the physical
properties of water
 Xylem sap rises against gravity, without the help of any mechanical pump, to
reach heights of more than 100 m in the tallest trees
 Sap flow is solar powered!
Transpirational pull and the ascent of xylem sap

-100 Mpa)
 Water lost to atmosphere is replaced by water from mesophyll, which is
replaced by water from the xylem stream
 For the most part, xylem sap is pulled upward by the leaves themselves

Transpiration provides the pull, and the cohesion of water due to hydrogen
bonding transmits the upward pull along the entire length of the xylem to
the roots
Transpiration-cohesion-adhesion mechanism
 Cohesion allows chains of water molecules to stay intact
 Adhesion to xylem vessel walls fights gravity
 Breaks in chain called cavitation can occur due to drought or freezing
Guard cell mediate the photosynthesis-transpiration compromise
 A leaf may transpire more than its weight in water each day
 To keep the leaf from wilting, flows in xylem vessels may reach 75 cm/min
 Guard cells, by
controlling the size
of stomata, help balance
the plant’s need to
conserve water with
its requirements for
photosynthesis
The compromise…
 Carbon dioxide diffuses in and oxygen and water vapor diffuses out of the
leaf via the stomata
 About 90% of the water that a plant loses escapes through stomata, though
these pores account for only 1 - 2 % of the external leaf surface
Water use efficiency
 One gauge of how efficiently a plant uses water is the transpiration-tophotosynthesis ratio, or water use efficiency the amount of water lost per gram
of CO2 assimilated into organic molecules by photosynthesis
 For many plant species, this ratio is about 600:1
 However, for corn and other C4 plants it is 300:1 or less
Transpiration
 The transpiration stream also assists in the delivery of minerals and other
substances from roots to the shoots and leaves
 Transpiration also results in evaporative cooling, which can lower the
temperature of a leaf by as much as 10-15 oC compared with the surrounding
air
Transpiration rates
 Vary with humidity, temperature, wind speed, leaf shape
 Many of which can influence boundary layer thickness around leaves
Wilting
 When transpiration exceeds the delivery of water by xylem, the leaves begin to
wilt as the cells lose turgor pressure
 Each stoma is flanked by a pair of guard cells which are suspended by other
epidermal cells over an air chamber, leading to the internal air space
Stomata
 Regulatory pores on undersurface of leaves, surrounded by guard cells,
control gas exchange
 Unique microfibril arrangement allows guard cells to buckle and pore to open
when cells are turgid
 Movement of K+ ions largely controls the degree of opening
Stomata
 In general, stomata are open during the day and closed at night to minimize
water loss when it is too dark for photosynthesis
 At least three cues contribute to stomatal opening at dawn
Stomatal opening
 First, blue-light receptors in the guard cells stimulate the activity of ATPpowered proton pumps in the plasma membrane, promoting the uptake of
K+
 A second stimulus is depletion of CO2 within air spaces of the leaf as
photosynthesis begins
 A third cue in stomatal opening is an internal clock located in the guard
cells
 The opening and closing cycle of the stomata is an example of a
circadian rhythm
Stomatal closure
 Various environmental stresses can cause stomata to close during the day
 Water deficiency causes guard cells to lose turgor
 Abscisic acid, a hormone produced by the mesophyll cells in response to
water deficiency, signals guard cells to close stomata
 High temperatures, by stimulating CO2 production by respiration, and
excessive transpiration may combine to cause stomata to close briefly
during mid-day
Factors that control stomata
 Low internal (often in high light) induces stomatal opening
 Low water availability induces stomatal closure
 Low light induces stomatal closure, because internal CO2 levels build up
 Abscisic acid (ABA) induces stomatal closure, role in drought resistance
Introduction
 The phloem transports the organic products of photosynthesis throughout the
plant via a process called translocation
 Phloem sap is an aqueous solution in which sugar, primarily the disaccharide
sucrose in most plants, is the most prevalent solute
 It may also contain minerals, amino acids, and hormones
From sources to sinks
 In contrast to the unidirectional flow of xylem sap from roots to leaves, the
direction that phloem sap travels is variable
 In general, sieve tubes carry food from a sugar source to a sugar sink
 A sugar source is a plant organ (especially mature leaves) in which sugar is
being produced by either photosynthesis or the breakdown of starch
 A sugar sink is an organ (such as growing roots, shoots, or fruit) that is a
net consumer or storer of sugar
Sources and sinks
 A storage organ, such as a tuber or a bulb, may be either a source or a sink,
depending on the season
 A sugar sink usually receives its sugar from the sources nearest to it
 One sieve tube in a vascular bundle may carry phloem sap in one direction
while sap in a different tube in the same bundle may flow in the opposite
direction
Pressure flow
 Phloem sap flows from source to sink at rates as great as 1 m/hr, faster than
can be accounted for by either diffusion or cytoplasmic streaming
 Phloem sap moves by bulk flow driven by pressure
 Higher levels of sugar at the source lowers the water potential and causes
water to flow into the tube
 Removal of sugar at the sink increases the water potential and causes
water to flow out of the tube
 The difference in hydrostatic pressure drives phloem sap from the source to
the sink
Phloem loading
 Sugar from mesophyll cells or other sources must be loaded into sieve-tube
members before it can be exported to sugar sinks
 In some species, sugar moves from mesophyll cells to sieve-tube members
via the symplast
 In other species, sucrose reaches sieve-tube members by a combination of
symplastic and apoplastic pathways
Phloem unloading
 Downstream, at the sink end of the sieve tube, phloem unloads its sucrose
 The mechanism of phloem unloading is highly variable and depends on
plant species and type of organ
 Because the concentration of free sugar in the sink is lower than in the
phloem, sugar molecules diffuse from the phloem into the sink tissues
 Water follows by osmosis


The pressure flow model explains why phloem sap always flows from sugar
source to sugar sink, regardless of their locations in the plant
The closer the aphid’s stylet is to a sugar source, the faster the sap will flow
out and the greater its sugar concentration