Overview
Transpiration is the
loss of water from a plant in the form of water vapour. Water is absorbed by
roots from the soil and transported as a liquid to the leaves via xylem. In the
leaves, small pores allow water to escape as a vapour. Of all the water absorbed
by plants, less than 5% remains in the plant for growth. This lesson will
explain why plants lose so much water, the path water takes through plants, how
plants might control for too much water loss to avoid stress conditions, and
how the environment plays a role in water loss from plants.
Objectives
At the completion of
the blog, you will be able to:
a. Define transpiration
and explain why it occurs in plants.
b. Follow the pathway
that water takes through plants from root uptake to evaporation at leaf cell
surfaces.
c. Describe how the
driving force for water movement and any resistances to its flow through the
plant are the two major components controlling rates of transpiration.
d. Describe how
environmental conditions alter rates of transpiration.
e. Explain how the
plant is able to alter rates of transpiration.
Introduction
Welcome to a blogpost
that will examine how water moves through plants. Plants lose gallons of water
every day through the process of transpiration, the evaporation of water from
plants primarily through pores in their leaves. Up to 99% of the water absorbed
by roots is lost via transpiration through plant leaves. How and why do they do
it? How do the plants avoid losing too much water? What environmental conditions
control water loss? The animation in the lesson will provide a visual tool for
you to understand these processes as well. The animation contains the text
below.
Did
you know that an acre of corn can transpire up to 400,000 gallons of water in one
growing season?
Because water molecules
stick together so well due to hydrogen bonding, water can be pulled up trees
100 meters tall (that’s over 300 feet or three times the height of most shade
trees!).
What is transpiration?
In actively growing plants,
water is continuously evaporating from the surface of leaf cells exposed to
air. This water is replaced by additional absorption of water from the soil.
Liquid water extends through the plant from the soil water to the leaf cell
surfaces where it is converted from a liquid into a gas through the process of
evaporation. The cohesive properties of water (hydrogen bonding between
adjacent water molecules) allow the column of water to be ‘pulled’ up through
the plant as water molecules are evaporating at the leaf surface. This process
has been termed the Cohesion Theory of
Sap Ascent in plants.
Why do plants transpire?
Evaporative cooling: As
water evaporates or converts from a liquid to a gas at the leaf cell and
atmosphere interface, energy is released. This exothermic process uses energy
to break the strong hydrogen bonds between liquid water molecules; the energy
used to do so is taken from the leaf and given to the water molecules that have
converted to highly energetic gas molecules. These gas molecules and their
associated energy are released into the atmosphere, cooling the plant.
Accessing
nutrients from the soil: The water that enters the root
contains dissolved nutrients vital to plant growth. It is thought that
transpiration enhances nutrient uptake into plants.
Carbon
dioxide entry: When a plant is transpiring, its stomata
are open, allowing gas exchange between the atmosphere and the leaf. Open
stomata allow water vapor to leave the leaf but also allow carbon dioxide (CO2)
to enter. Carbon dioxide is needed for
photosynthesis to operate. Unfortunately, much more water leaves the leaf than
CO2 enters for three reasons:
1. H2O
molecules are smaller than CO2 molecules and so they move to their
destination faster.
2. CO2 is only about 0.036% of the
atmosphere (and rising!) so the gradient for its entry into the plant is much
smaller than the gradient for H2O moving from a hydrated leaf into a
dry atmosphere.
3. CO2 has a
much longer distance to travel to reach its destination in the chloroplast from
the atmosphere compared to H2O which only has to move from the leaf
cell surface to the atmosphere. This disproportionate exchange of CO2
and H2O leads to a paradox. The larger the stomatal opening, the
easier it is for carbon dioxide to enter the leaf to drive photosynthesis;
however, this large opening will also allow the leaf to lose large quantities
of water and face the risk of dehydration or water-deficit stress. Plants that
are able to keep their stomata slightly open, will lose fewer water molecules
for every CO2 molecule that enters and thus will have greater water
use efficiency (water lost/CO2 gained). Plants with greater water
use efficiencies are better able to withstand periods when water in the soil is
low.
Water uptake
Although only less than
5% of the water taken up by roots remains in the plant, that water is vital for
plant structure and function. The water is important for driving biochemical
processes, but also it creates turgor so that the plant can stand without
bones.
How fast does water
move through plants? Transpiration rates depend on two major factors: 1) the
driving force for water movement from the soil to the atmosphere and 2) the
resistances to water movement in the plant.
Driving force
The driving force for
transpiration is the difference in water potential between the soil and the
atmosphere surrounding the plant. This difference creates a gradient, forcing
water to move toward areas with less water. The drier the air around the plant,
the greater the driving force is for water to move through the plant and the
faster the transpiration rate.
Resistances
There are three major
resistances to the movement of water out of a leaf: cuticle resistance, stomata
resistance and boundary layer resistance. These resistances slow water
movement. The greater any individual resistance is to water movement, the
slower the transpiration rate.
A simple equation
describing how these factors alter transpiration is:
Transpiration = [Water potential (leaf)] – [Water potential
(atmosphere)/Resistance
The units for this
equation are moles of water lost per leaf area per time (mole/cm2/s). This
equation makes predicting rates of transpiration easy. For example, any time
the numerator (the value for the driving force) is increased, the rate of
transpiration becomes faster and vice versa. Similarly, if the denominator (the
value for resistance) increases, this means there is greater resistance and
thus, slower transpiration.
Major Plant Highlights
Root Detail: The major
path for water movement into plants is from soil to roots. Water enters near
the tip of a growing root, the same region where root hairs grow. The surface
of the root hairs needs to be in close contact with the soil to access soil
water. Water diffuses into the root, where it can take at least three different
pathways to eventually reach the xylem, the conduit located at the interior of
the root that carries the soil water to the leaves. View the next level of this
animation to see the possible pathways that water can take across a root.
What path does water
take to reach the leaf from the root hair? Once water has entered a root hair,
it must move across the cortex and endodermis before it reaches the xylem.
Water will take the path of least resistance through a root to reach the xylem.
Water can move across the root via three different pathways. One path is the
apoplastic path where the water molecule stays between cells in the cell wall
region, never crossing membranes or entering a cell. The other two routes,
called cellular pathways, require the water molecule to actually move across a
membrane. The first cellular pathway is the transmembrane path where water
moves from cell to cell across membranes; it will leave one cell by traversing
its membrane and will re-enter another cell by crossing its membrane. The
second cellular path is the symplastic path which takes the water molecule from
cell to cell using the intercellular connections called the plasmodesmata which
are membrane connections between adjacent cells. Regardless of the pathway,
once the water molecule has traversed the cortex, it must now cross the
endodermis. The endodermis is a layer of cells with a waxy inlay or mortar
called the Casparian strip that stops water movement between cells. At this
point, water is forced to move through the membranes of endodermal cells,
creating a sieving effect. Once in the endodermal cells, the water freely
enters the xylem cells where it joins the fast moving column of water or
transpiration stream, headed to the leaves.
Xylem
Details: The xylem is probably the longest part of the
pathway that water takes on its way to the leaves of a plant. It is also the
path of least resistance, with about a billion times less resistance than cell
to cell transport of water. Xylem cells are called tracheids (cells with
narrower diameters) or vessels (cells with wider diameters). Their cell walls
contain cellulose and lignin making them extremely rigid. Xylem cells contain
no membranes and are considered dead. These cells overlap to create a series of
pathways that water can take as it heads to the leaves. There is no single
column of xylem cells carrying water.
Cavitation:
Cavitation is the filling of a xylem vessel or tracheid with air. It is also
known as an ‘embolism’ or ‘air-lock’. Remember that during transpiration, the
column of water is being pulled out of the plant by evaporation at the leaf
cell surface. When this ‘pulling’ of water out of the plant becomes greater
than the ability of the water molecules to stay together, the column of water
will break. Using sound-sensing equipment, one can actually hear a ‘click’ when
the water molecules split from one another. Unique structural characteristics
help the plant contain the air bubble so that it does not totally disrupt water
movement up the plant.
Plants are particularly
sensitive to cavitation during the hottest part of the day when there is not
enough water available from the soil to keep up with the demand for water while
it is evaporating off the leaf surface. Cavitation also occurs under freezing
conditions. Because the solubility of gas in ice is very low, gas comes out of
solution when the xylem sap freezes. Freezing of xylem sap is a problem in the
spring when the ice thaws, leaving a bubble in a xylem vessel. These bubbles
can block water transport and cause water deficit in leaves.
Plants avoid cavitation
or minimize its damage through several mechanisms:
1.
Xylem cells possess pits or tiny holes that allow
liquid water transport, but do not allow the gas bubble to escape; this
structural characteristic helps keep the gas bubble in one cell, so the other
xylem cells can continue to transport water up the plant.
2) Water will detour around any xylem cell containing an air bubble
through the pits as well.
3) The gas bubble will re-dissolve into liquid water when the pulling
of water through the xylem is reduced, such as during the night when water is
not being pulled out of the leaf via transpiration because the stomata are
closed.
4) Xylem cells with narrower diameters (tracheids) compared to those
with wider diameters (vessels) avoid cavitation because the column of water in
a cell with a narrow diameter is better able to resist bubble formation or
rupture.
Plants are most
susceptible to cavitation when transpiration rates are extremely high.
Stomata
Details: The stomata are the primary control mechanisms that
plants use to reduce water loss and they are able to do so quickly. Stomata are
sensitive to the environmental cues that trigger the stomata to open or close.
The major role of stomata is to allow carbon dioxide entry to drive
photosynthesis and at the same time allow the exit of water as it evaporates,
cooling the leaf. Two specialized cells called ‘guard cells’ make up each stoma
(stoma is singular for stomata). Plants have many stomata (up to 400 per mm2)
on their leaf surfaces and they are usually on the lower surface to minimize
water loss.
Webster’s Dictionary
says a stoma (singular form of stomata) is a small, simple opening.
How
do stomata open? Stomata sense environmental cues, like
light, to open. These cues start a series of reactions that cause their guard
cells to fill with water. Let’s follow a scenario where the sun is rising and a
cotton plant is signaled to open its stomata.
1. Signal received: The blue light at dawn is
the signal that is recognized by a receptor on the guard cell.
2. The receptor signals
the H+-ATPases on the guard cell’s plasma membrane to start pumping protons
(H+) out of the guard cell. This loss of positive charge creates a negative
charge in the cell.
3. Potassium ions (K+)
enter the guard cell through channels in the membrane, moving toward its more
negative interior.
4. As the potassium
ions accumulate in the guard cell, the solute potential is lowered.
5. A lower solute
potential attracts water to enter the cell.
6. As water enters the
guard cell, its hydrostatic pressure increases.
7. The pressure causes
the shape of the guard cells to change and a pore is formed, allowing gas
exchange.
Side View of Stomata
Environmental cues that
affect stomata opening and closing are light, water, temperature, and the
concentration of CO2 within the leaf. Stomata will open in the light
and close in the dark. However, stomata can close in the middle of the day if
water is limiting, CO2 accumulates in the leaf, or the temperature
is too hot.
Stomata Diagram |
If the plant lacks
water, stomata will close because there will not be enough water to create
pressure in the guard cells for stomatal opening; this response helps the plant
conserve water.
If the leaf’s internal
concentration of CO2 increases, the stomata are signaled to close
because respiration is releasing more CO2 than photosynthesis is
using. There is no need to keep the stomata open and lose water if
photosynthesis is not functioning. Alternatively, if the leaf’s CO2 concentration
is low, the stomata will stay open to continue fueling photosynthesis.
High temperatures will
also signal stomata to close. High temperatures will increase the water loss
from the leaf. With less water available, guard cells can become flaccid and
close. Another effect of high temperatures is that respiration rates rise above
photosynthesis rates causing an increase of CO2 in the leaves; high
internal CO2 will cause stomata to close as well. Remember that some
plants may open their stomata under high temperatures so that transpiration
will cool the leaves.
Why do plants waste all
this water!? Transpiration helps the plant by providing evaporative cooling,
nutrient uptake, and carbon dioxide entry.
Top View of Stomata
Open
Stomata: When stomata are signaled to open, potassium ions
(K+) enter the guard cells. This causes water to enter down its water potential
gradient, creating a hydrostatic pressure in the guard cell that changes the
shape of the stoma. Guard cells expand on the outer edges of the stoma, but not
on the inner side, resulting in kidney-shaped cells and an opening or pore
between the two guard cells for gas exchange.
The shape taken by the
guard cells is dependent on cellulose microfibrils that fan out radially from
the pore, somewhat similar to radial tires. The cellulose microfibrils are
rigid and do not stretch when water has entered the cell. The cell walls
surrounding the stomatal opening are thickened, preventing that side of the
guard cell from expanding. Therefore, when pressure in the cell increases due
to water entry, guard cell does not widen, but rather the outer edge stretches
disproportionately more than the inner edge. This unequal stretching allows the
pore to form between the two guard cells.
Cellulose microfibrils
keep the guard cells from expanding into sphere-shaped cells because the
fibrils are rigid rings around the guard cell, restricting cell expansion other
than the outer edge.
Closed
Stomata: Stomata must be open for the plant to
photosynthesize; however, open stomata present a risk of losing too much water
through transpiration. Stomata close when the guard cells lose water and become
flaccid. This occurs because potassium ions move back out of the guard cell,
followed by water that lowers the pressure in the guard cell.
Many plant leaves have
stomata only on their lower surface to help avoid water loss.
Factors Affecting Rates of
Transpiration
Plant
Parameters: These plant parameters help plants
control rates of transpiration by serving as forms of resistance to water
movement out of the plant.
Stomata:
Stomata are pores in the leaf that allow gas exchange where water vapor leaves
the plant and carbon dioxide enters. Special cells called guard cells control
each pore’s opening or closing. When stomata are open, transpiration rates
increase; when they are closed, transpiration rates decrease.
Boundary
layer: The boundary layer is a thin layer of still air
hugging the surface of the leaf. This layer of air is not moving. For
transpiration to occur, water vapor leaving the stomata must diffuse through
this motionless layer to reach the atmosphere where the water vapor will be
removed by moving air. The larger the boundary layer, the slower the rates of
transpiration.
Plants can alter the
size of their boundary layers around leaves through a variety of structural
features. Leaves that possess many hairs or pubescence will have larger
boundary layers; the hairs serve as mini-wind breaks by increasing the layer of
still air around the leaf surface and slowing transpiration rates. Some plants
possess stomata that are sunken into the leaf surface, dramatically increasing
the boundary layer and slowing transpiration. Boundary layers increase as leaf
size increases, reducing rates of transpiration as well. For example, plants
from desert climates often have small leaves so that their small boundary
layers will help cool the leaf with higher rates of transpiration.
Did
you know that hairs on a leaf surface increase the boundary layer and thus,
reduce rates of transpiration?
Cuticle:
The cuticle is the waxy layer present on all above-ground tissue of a plant and
serves as a barrier to water movement out of a leaf. Because the cuticle is made of wax, it is
very hydrophobic or ‘water- repelling’; therefore, water does not move through
it very easily. The thicker the cuticle layer on a leaf surface, the slower the
transpiration rate.
Cuticle thickness
varies widely among plant species. In general, plants from hot, dry climates
have thicker cuticles than plants from cool, moist climates. In addition,
leaves that develop under direct sunlight will have much thicker cuticles than
leaves that develop under shade conditions.
Sun leaves have much
thicker cuticles than shade leaves causing slower rates of transpiration.
Environmental Conditions – Some environmental conditions create the driving
force for movement of water out of the plant. Others alter the plant’s ability
to control water loss.
Relative
humidity: Relative humidity (RH) is the amount of water vapor
in the air compared to the amount of water vapor that air could hold at a given
temperature. A hydrated leaf would have a RH near 100%, just as the atmosphere
on a rainy day would have. Any reduction in water in the atmosphere creates a
gradient for water to move from the leaf to the atmosphere. The lower the RH,
the less moist the atmosphere and thus, the greater the driving force for
transpiration. When RH is high, the atmosphere contains more moisture, reducing
the driving force for transpiration.
The drier the
atmosphere, the larger the driving force for water movement out of the plant,
increasing rates of transpiration.
Temperature:
Temperature greatly influences the magnitude of the driving force for water
movement out of a plant. As temperature increases, the water holding capacity
of that air increases sharply. The amount of water does not change, just the
ability of that air to hold water. Because warmer air can hold more water, its
relative humidity is less than the same air sample at a lower temperature, or
it is ‘drier air’. Because cooler air holds less water, its relative humidity
increases or it is ‘moister air’. Therefore, warmer air will increase the driving
force for transpiration and cooler air will decrease the driving force for
transpiration.
Warmer air holds more
water, creating a larger driving force for water movement out of the plant,
increasing rates of transpiration.
Light:
Stomata are triggered to open in the light so that carbon dioxide is available
for the light-dependent process of photosynthesis. Stomata are closed in the
dark in most plants. Very low levels of light at dawn can cause stomata to open
so they can access carbon dioxide for photosynthesis as soon as the sun hits
their leaves. Stomata are most sensitive to blue light, the light predominating
at sunrise.
Light levels as low as
one thousandth of the sun can cause stomata to open.
Wind:
Wind
can alter rates of transpiration by removing the boundary layer, that still
layer of water vapor hugging the surface of leaves. Wind increases the movement
of water from the leaf surface when it reduces the boundary layer, because the
path for water to reach the atmosphere is shorter.
Windier conditions
increase transpiration because the leaf’s boundary layer is smaller.
Summary
Transpiration is the
evaporation of water from the surface of leaf cells in actively growing plants.
This water is replaced by additional absorption of water from the soil leading
to a continuous column of water in the plant's xylem. The process of
transpiration provides the plant with evaporative cooling, nutrients, carbon
dioxide entry and water to provide plant structure. Rates of transpiration
depend on the water potential gradient from the soil to the atmosphere and the
resistances to its movement through the plant. Water enters the root and
travels through the cortex and endodermal layers of cells to reach the xylem
where water ascends to the leaf where, if not used in the plant, evaporates. If
water loss is greater than water uptake, air bubbles can form in the xylem.
Plants reduce water loss by closing their stomata, developing thick cuticles,
or by possessing leaf hairs to increase the boundary layer. Stomata are quick
to respond to environmental cues to protect the plant from losing too much
water, but still allowing in enough carbon dioxide to drive photosynthesis.
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