THEME 2: WATER UPTAKE AND TRANSPORT

THROUGH PLANTS
WATER IN THE SOIL
After heavy rain, water fills every space between soil particles. A portion of this water,
named gravitational water or ground water, percolates downward by gravity, eventually
becoming unavailable to plant roots. The moisture-holding capacity of soil is called the
field capacity, but when the soil dries, water becomes progressively less available to
plants. Furthermore, the permanent wilting point depends on the soil and the plant species.
When the water is held on the surface of soil particles we call it
hygroscopic water, but the one that remains available is the capillary
water one. There is also air or gravitational water that helps draining
rapidly down the water table in all except the most compact of soils.

SOIL WATER STATUS QUANTIFICATION

- Soil water content: can be measured on a volumetric basis by time domain
reflectometry, or gravimetrically (by weighing a sample before and after oven
drying at 105oC)
- Soil water potential: can be measured with tensiometers (yM) or psicrometers. It
describes the energy status of soil water.
Either way, values would refer only to small portion of soil sampled. Predawn leaf water
potential is a mostly reliable estimate of water availability in the portion of soil explored
by fine roots (rhizosphere).

y(root) < y(soil)

The pattern of this relationship is like that obtained for the relationship between soil water
content and predawn leaf water potential, which further supports the reliability of
predawn leaf water potential to quantify soil water availability to plants.
Exceptions: nocturnal transpiration, extensive cavitation, short night, high xylem
resistance to flow, …
FACTORS AFFECTING SOIL WATER AVAILABILITY
Based in the reaching and absorbing water process:
- Concentration and composition of soil solution: excessive salinity is seldom a
problem in forested areas. However, it may become a problem in forest plantations
established on previously cultivated and irrigated soils.
- Soil aeration: oxygen deficiency in flooded soils may reduce the capacity of water
channel proteins to pass water through root membranes.
- Soil temperature: low soil temperatures reduce water uptake in two ways:
o Directly: by decreasing the permeability of roots to water.
o Indirectly: by increasing the viscosity of water, which slows its movement
through both soil and roots.
o Extent and efficiency of root systems: differences between species and
alterations due to environment factors.

WATER ABSORPTIONS BY ROOTS: REACHING WATER

Tropism: orientated plant growth in response to a perceived directional stimulus from
light, gravity, or touch. Root growth shows:
- Positive gravitropism (growing downward into the soil)
- Thighmotropism (growing over rocks surface to override the obstacle)
- Chemotropism (root proliferation in fertilized patches)
- Hydrotropism (root proliferation in wet soil patches or inside pipes.)
The larger the volume of the soil explored by roots, the larger amount of water available
to the plant. Water moves in the soil by bulk flow, capillarity, and diffusion. Water moves
slowly in most soils. Hence, roots must grow in search of water.

WATER ABSORPTION BY PLANTS

Plants take water from the groundwater that can be shallow or deep. Typically, root
density is highest near soil surface (shallow), because in the topsoil there are much more
mineral nutrients due to the dead organic material mostly been in the surface. Otherwise,
if there is not enough shallow water, they uptake the deep one.
Shallow water and deep water differ in their isotropic composition. 1H216O is the most
abundant water molecules, but there are more like: 1H218O, 2H216O, 2H218O.
Isotopic composition is quantified by mass spectrometry:
[ "#𝑂] (𝑟𝑎𝑟𝑒 𝑖𝑠𝑜𝑡𝑜𝑝𝑒)
R=
[ "$𝑂 ](𝑐𝑜𝑚𝑚𝑜𝑛𝑖𝑠𝑜𝑡𝑜𝑝𝑒)
Isotopic composition increases with the abundance of the rare isotope:

δ = 10007R %&'()* /R %+&,-&.- − 1:

Fractionation changes isotopic composition, it may occur during condensation or

biochemistry, leading to changes in the isotopic composition of rainwater through the
year and changes in 𝛿 18 O and d2H inside the plant.

WATER ABSORPTION BY ROOTS:

Water enters the root mainly near the root tip where:
- Specialized epidermic cells named trichoblasts (root hairs) significantly increase
the absorptive root surface.
- Cork cambium is absent and therefore the outer layer of tissue is not suberized as
occurs in older portions of the root.
Water enters the root mainly by bulk form, due to the existence of a water potential
gradient between the root and the soil. Most water transport is passive, but active uptake
of water may also occur:
- If the water potential gradient appears because of the water loss from leaves by
transpiration, then water transport is said to be passive (water loss by
transpiration).
- If the water potential gradient appears because of solute accumulation in root cells,
then water transport is said to be active (solute accumulation), because metabolic
energy is needed to accumulate solutes.
Active absorption of water may lead to root pressure and guttation. Water exits the leaves
through hidathodes, which are specialized structures typically placed at the end of leaf
veins, close to the leaf margin.
Guttation: water exudation from leaves or shoots due to the generation of positive
hydrostatic pressure in xylem vessel. This phenomenon occurs most frequently after
warm nights, when soil water is abundant, and the air is humid.

HYDRAULIC REDISTRIBUTION
Using stable hydrogen isotope analysis, showed that water absorbed by deep roots of
sugar maple trees was released into shallow soil layers where it was absorbed by
understory plants.
Advantages to trees:
- Water moved through the night to shallow soil layers can contribute to daily
transpiration.
- The release of water in the vicinity of roots may have benefits if increased nutrient
from otherwise dry soil.
- Some of the shrubs and herbaceous species favored by the water lift by trees may
improve soil fertility (plant species able to establish symbiosis with nitrogen-
fixing soil bacteria).
The isotopic signature of rainfall and shallow soil water differ from that deep water.
Dawson found that close to the stem of sugar maple trees soil water showed an isotopic
signature characteristic of deep water. This hydraulic redistribution is often relevant in
the tropics during the dry season.
WATER TRANSPORT IN ROOT
The water moves evenly because it enters mainly by the primary body bidirectionally
(homogeneous redistribution and absorption):
- In moist soil: root cell membranes are the greatest resistance to water flow through
the root.
- In dry soil: contact between roots and soil becomes the greatest resistance to water
movement from soil to leaves.
Theoretical pathways:
There are 3, but they do not exclude each other, they are complementary
- Apoplastic pathway (easiest): crossing no membranes, cells with primary wall
with highly hydrophobic cell walls and intercellular spaces. Until they Casparian
strip/endodermis because you must cross it obligatory
- Symplastic pathwaty: water enters on first cell and then moves from cell to cell
via the plasmodesmata.
- Transcellular pathway (hardest): water must cross a high number of cell
membranes. Remember that the inner layer of biological membranes is
hydrophobic. Slow movement.
There are also aquaporins which are membrane proteins that facilitate the transport of
water across the membrane. The hydrophobic nature of the Casparian Strip forces
apoplastic water to enter these cells to reach the tracheary
elements of the xylem to be transported to the trunk and leaves.
THE COHESION-TENSION THEORY AND THE ASCENT OF SAP
Pressure-driven bulk flow is responsible for long-distance transport of
water in the xylem. The pressure gradient needed to move water results
from water loss due to transpiration, which decreases y(leaf). The
energy needed to pull up water against gravity along a path that can
exceed 100 m in the tallest trees comes from the sun: solar radiation,
increases the temperature of both the leaf and the surrounding air which
favors water evaporation inside the leaf water vapor diffusion from the
leaf to the atmosphere.
Water inside the tracheary elements of the xylem is under tension. The
cohesive forces between water molecules would allow water to remain
liquid, according to the cohesion-tension theory.
The results of the next graph support the cohesion-tension theory.

At dawn, the slope of the relationship shows a rate

of decrease of y = 0.01 MPa/m which is the strict
gradient needed to keep water against gravity. At
noon, y must be more negative to overcome
frictional resistances and move water

WATER TRANSPORT THROUGH THE XYLEM

Only a fraction of the xylem’s surface is used for sap transport (Heartwood // Sapwood,
bit not only) neither the stem diameter nor the sum of vessel/tracheid surface sections
reflect the ability of the stem transport sap.
The tracheary elements (vessels/tracheids) can lose functionality by means of cavitation
and embolism (both of then can occur consecutively in transpiring plants):
- Cavitation: the formation of small vapor-filled cavities (bubbles) within a liquid
at low-pressure regions. In plants, cavitation occurs in the xylem spa.
 - Embolism: the sudden obstruction of a tracheary element by the collapse of small
vapor-pressure bubbles to forma a largest, air-filled bubble. This can be repaired
by refilling embolized vessels/tracheids.
This cavitation can start a freeze/thaw cycle or drop the water potential. To solve the
second problem, there is a water uptake/loss imbalance that increases transpiration and
reduce the soil water availability. The consequences of this problem can be vascular
diseases such as Dutch elm disease, pine wilt disease (pinewood nematode).
The impact of embolism can be, measured by destructive and non-destructive techniques.
The first one consists in the measuring of the hydraulic conductance of a stem in its native
state and after removing embolism with pressurized water, and the other one is by
ultrasonic emissions from the stem.
Hydraulic conductance can be measured by placing a portion of the stem in a tubing
system. A low-pressure water solution is used to measure water flow through the stem .
this value is then corrected by the pressure applied to move water, to calculate the initial
or native hydraulic conductance of the stem (Ki). Once the initial flow is measured, a
high-pressure water solution is applied to remove embolism. Maximum hydraulic
conductance (KM) is measured afterwards.
Hydraulic conductivity is calculated by multiplying the
hydraulic conductance by the length of the stem. The
percentage loss of hydraulic conductance is calculated as:
𝐾/ −𝐾0
𝑃𝐿𝐶(%) = 100
𝐾/
The vulnerability curves are the plot of the percentage loss of
hydraulic conductance versus xylem water potential. When
they reach the 100% is the most vulnerable to embolism
situation, in the other hand the more negative the water potential, more resistant to
embolism. This are the reasons why there is a tendency of species living in more xeric
environments to show higher resistance to embolism.
The ascent of sap follows a curvilinear pattern rather than a straight one, as the result of:
- Water in the xylem conducts/vessels moves according to the vertical component
and from one vessel to another through the pits of the neighbor.
- The vessels/tracheids form an angle that is named spiral grain and is a defect
economically for the wood because it has some tension and devaluates its price.
The water movement is an slow process and the amount of water moved variates due to
the different radius of the vessels/tracheids.
The water stored in the stem may contribute to the ascent of sap. Water uptake in many
trees lags behind transpirational water loss by about 2 hours. The ability of the stem to
store water can be quantified by measuring the change in the column of water stored as
water potential decreases (hydraulic capacitance, C)
∆𝑉
𝐶(𝑚1 𝑀/𝑃𝑎) =
∆𝜑
Changes in the amount of water stored lead to reversible shrinking and swelling of the
stems (depends of the elasticity of the cell/tissue), which can be detected by attaching
linear variable displacement transducers (LVDT) to the stem.
Graphical output obtained from LVDT sensors: The plant well-watered (yellow) during
the daytime does not lose a big amount
of water, that is why shrinking is less
present than in the blue one.
Another difference is the density of the
wood, the denser the wood, the less
water store. Also, the less wood
density, more vulnerable to cavitation
and a higher water potential value.
Pinus stores and lose more water than
Quercus.

During the nighttime we have swelling (the stem

rehydrates fast and stabilize the inner temperature) and
during daytime we have shrinking (as temperatures
increases, gradients increase, and the stem loses water).
The dehydration affects equally y, yP and yS, but yP and
yS decrease less than y, because is the addition of both so
it decreases more, till 0 than means it is a full turgor.