Transpiration

Transpiration is the process of water

movement through a plant and its
evaporation from aerial parts, such as
leaves, stems and flowers. Water is
necessary for plants but only a small
amount of water taken up by the roots is
used for growth and metabolism. The
remaining 97–99.5% is lost by
transpiration and guttation.[1] Leaf
surfaces are dotted with pores called
stomata (singular "stoma"), and in most
plants they are more numerous on the
undersides of the foliage. The stomata
are bordered by guard cells and their
stomatal accessory cells (together
known as stomatal complex) that open
and close the pore.[2] Transpiration
occurs through the stomatal apertures,
and can be thought of as a necessary
"cost" associated with the opening of the
stomata to allow the diffusion of carbon
dioxide gas from the air for
photosynthesis. Transpiration also cools
plants, changes osmotic pressure of
cells, and enables mass flow of mineral
nutrients and water from roots to shoots.
Two major factors influence the rate of
water flow from the soil to the roots: the
hydraulic conductivity of the soil and the
magnitude of the pressure gradient
through the soil. Both of these factors
influence the rate of bulk flow of water
moving from the roots to the stomatal
pores in the leaves via the xylem.[3]

Overview of transpiration:
1. Water is passively transported into the roots
and then into the xylem.
2. The forces of cohesion and adhesion cause
the water molecules to form a column in the
xylem.
3. Water moves from the xylem into the
mesophyll cells, evaporates from their
surfaces and leaves the plant by diffusion
 through the stomata

Transpiration of Water in Xylem

Stoma in a tomato leaf shown via colorized

scanning electron microscope
The clouds in this image of the Amazon Rainforest

are a result of evapotranspiration.

Mass flow of liquid water from the roots

to the leaves is driven in part by capillary
action, but primarily driven by water
potential differences. If the water
potential in the ambient air is lower than
the water potential in the leaf airspace of
the stomatal pore, water vapor will travel
down the gradient and move from the
leaf airspace to the atmosphere. This
movement lowers the water potential in
the leaf airspace and causes evaporation
of liquid water from the mesophyll cell
walls. This evaporation increases the
tension on the water menisci in the cell
walls and decrease their radius and thus
the tension that is exerted on the water in
the cells. Because of the cohesive
properties of water, the tension travels
through the leaf cells to the leaf and
stem xylem where a momentary negative
pressure is created as water is pulled up
the xylem from the roots.[4] As
evaporation occurs at the leaf surface,
the properties of adhesion and cohesion
work in tandem to pull water molecules
from the roots, through xylem tissue, and
out of the plant through stomata.[5] In
taller plants and trees, the force of gravity
can only be overcome by the decrease in
hydrostatic (water) pressure in the upper
parts of the plants due to the diffusion of
water out of stomata into the
atmosphere. Water is absorbed at the
roots by osmosis, and any dissolved
mineral nutrients travel with it through
the xylem.

The cohesion-tension theory explains

how leaves pull water through the xylem.
Water molecules stick together, or exhibit
cohesion. As a water molecule
evaporates from the surface of the leaf, it
pulls on the adjacent water molecule,
creating a continuous flow of water
through the plant.[6]

Regulation
Plants regulate the rate of transpiration
by controlling the size of the stomatal
apertures. The rate of transpiration is
also influenced by the evaporative
demand of the atmosphere surrounding
the leaf such as boundary layer
conductance, humidity, temperature,
wind and incident sunlight. Along with
above ground factors, soil temperature
and moisture can influence stomatal
opening,[7] and thus transpiration rate.
The amount of water lost by a plant also
depends on its size and the amount of
water absorbed at the roots.
Transpiration accounts for most of the
water loss by a plant by the leaves and
young stems. Transpiration serves to
evaporatively cool plants, as the
evaporating water carries away heat
energy due to its large latent heat of
vaporization of 2260 kJ per litre.
 Feature Effect on transpiration

More leaves (or spines, or other photosynthesizing organs) means a bigger

Number of
surface area and more stomata for gaseous exchange. This will result in
leaves
greater water loss.

Number of
More stomata will provide more pores for transpiration.
stomata

Size of the A leaf with a bigger surface area will transpire faster than a leaf with a smaller
leaf surface area.

A waxy cuticle is relatively impermeable to water and water vapour and

reduces evaporation from the plant surface except via the stomata. A
reflective cuticle will reduce solar heating and temperature rise of the leaf,
Presence of helping to reduce the rate of evaporation. Tiny hair-like structures called
plant cuticle trichomes on the surface of leaves also can inhibit water loss by creating a
high humidity environment at the surface of leaves. These are some examples
of the adaptations of plants for conservation of water that may be found on
many xerophytes.

The rate of transpiration is controlled by stomatal aperture, and these small

pores open especially for photosynthesis. While there are exceptions to this
Light supply
(such as night or CAM photosynthesis), in general a light supply will
encourage open stomata.

Temperature affects the rate in two ways:

1) An increased rate of evaporation due to a temperature rise will hasten the

Temperature loss of water.
2) Decreased relative humidity outside the leaf will increase the water
potential gradient.

Relative Drier surroundings gives a steeper water potential gradient, and so increases
humidity the rate of transpiration.

Wind In still air, water lost due to transpiration can accumulate in the form of vapor
close to the leaf surface. This will reduce the rate of water loss, as the water
potential gradient from inside to outside of the leaf is then slightly less. Wind
blows away much of this water vapor near the leaf surface, making the
potential gradient steeper and speeding up the diffusion of water molecules
into the surrounding air. Even in wind, though, there may be some
accumulation of water vapor in a thin boundary layer of slower moving air next
to the leaf surface. The stronger the wind, the thinner this layer will tend to be,
and the steeper the water potential gradient.
Water Water stress caused by restricted water supply from the soil may result in
supply stomatal closure and reduce the rates of transpiration.

The effect of temperature on the

transpiration rate of plants.

The effect of wind velocity on the

transpiration rate of plants.
 The effect of humidity on the
transpiration rate of plants.

Some xerophytes will reduce the surface of their

leaves during water deficiencies (left). If
temperatures are cool enough and water levels are
adequate the leaves expand again (right).

During a growing season, a leaf will

transpire many times more water than its
own weight. An acre of corn gives off
about 3,000–4,000 gallons (11,400–
15,100 liters) of water each day, and a
large oak tree can transpire
40,000 gallons (151,000 liters) per year.
The transpiration ratio is the ratio of the
mass of water transpired to the mass of
dry matter produced; the transpiration
ratio of crops tends to fall between 200
and 1000 (i.e., crop plants transpire 200
to 1000 kg of water for every kg of dry
matter produced).[8]

Transpiration rates of plants can be

measured by a number of techniques,
including potometers, lysimeters,
porometers, photosynthesis systems and
thermometric sap flow sensors. Isotope
measurements indicate transpiration is
the larger component of
evapotranspiration.[9] Recent evidence
from a global study[10] of water stable
isotopes shows that transpired water is
isotopically different from groundwater
and streams. This suggests that soil
water is not as well mixed as widely
assumed.[11]

Desert plants have specially adapted

structures, such as thick cuticles,
reduced leaf areas, sunken stomata and
hairs to reduce transpiration and
conserve water. Many cacti conduct
photosynthesis in succulent stems,
rather than leaves, so the surface area of
the shoot is very low. Many desert plants
have a special type of photosynthesis,
termed crassulacean acid metabolism or
CAM photosynthesis, in which the
stomata are closed during the day and
open at night when transpiration will be
lower.

Cavitation
To maintain the pressure gradient
necessary for a plant to remain healthy
they must continuously uptake water
with their roots. They need to be able to
meet the demands of water lost due to
transpiration. If a plant is incapable of
bringing in enough water to remain in
equilibrium with transpiration an event
known as cavitation occurs.[12] Cavitation
is when the plant cannot supply its xylem
with adequate water so instead of being
filled with water the xylem begins to be
filled with water vapor. These particles of
water vapor come together and form
blockages within the xylem of the plant.
This prevents the plant from being able
to transport water throughout its
vascular system.[13] There is no apparent
pattern of where cavitation occurs
throughout the plant's xylem. If not
effectively taken care of, cavitation can
cause a plant to reach its permanent
wilting point, and die. Therefore, the plant
must have a method by which to remove
this cavitation blockage, or it must create
a new connection of vascular tissue
throughout the plant.[14] The plant does
this by closing its stomates overnight,
which halts the flow of transpiration. This
then allows for the roots to generate over
0.05 mPa of pressure, and that is
capable of destroying the blockage and
refilling the xylem with water,
reconnecting the vascular system. If a
plant is unable to generate enough
pressure to eradicate the blockage it
must prevent the blockage from
spreading with the use of pit pears and
then create new xylem that can re-
connect the vascular system of the
plant.[15]

Scientists have begun using magnetic

resonance imaging (MRI) to monitor the
internal status of the xylem during
transpiration, in a non invasive manner.
This method of imaging allows for
scientists to visualize the movement of
water throughout the entirety of the
plant. It also is capable of viewing what
phase the water is in while in the xylem,
which makes it possible to visualize
cavitation events. Scientists were able to
see that over the course of 20 hours of
sunlight more than 10 xylem vessels
began filling with gas particles becoming
cavitated. MRI technology also made it
possible to view the process by which
these xylem structures are repaired in the
plant. After three hours in darkness it
was seen that the vascular tissue was
resupplied with liquid water. This was
possible because in darkness the
stomates of the plant are closed and
transpiration no longer occurs. When
transpiration is halted the cavitation
bubbles are destroyed by the pressure
generated by the roots. These
observations suggest that MRIs are
capable of monitoring the functional
status of xylem and allows scientists to
view cavitation events for the first
time.[14]
See also
Antitranspirant – a substance to prevent
transpiration
Canopy conductance
Ecohydrology
Eddy covariance flux (aka eddy correlation,
eddy flux)
Hydrology (agriculture)
Latent heat flux
Perspiration
Soil plant atmosphere continuum
Stomatal conductance
Transpiration stream
Turgor pressure
Water Evaluation And Planning system
(WEAP)
 Wikisource has the text of the 1905 New
International Encyclopedia article
"Transpiration".

References
1. Sinha, Rajiv Kumar (2004-01-01).
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4. Freeman, Scott (2014). Biological
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(2019). Campbell essential biology.
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transpiration: a plot scale
 manipulation in a young Scots pine
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Fawcett, Peter J. (3 April 2013).
"Terrestrial water fluxes dominated
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 Bibcode:2013Natur.496..347J .
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10. Evaristo, Jaivime; Jasechko, Scott;
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"Global separation of plant
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"Stomatal Closure, Basal Leaf
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 Plant Physiology. 174 (2): 764–775.
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External links
 USGS The Water Cycle:
Evapotranspiration

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