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Transpiration is the process of
water Water (chemical formula ) is an inorganic, transparent, tasteless, odorless, and nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living organisms (in which it acts as a ...
movement through a
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...
and its
evaporation Evaporation is a type of vaporization that occurs on the surface of a liquid as it changes into the gas phase. High concentration of the evaporating substance in the surrounding gas significantly slows down evaporation, such as when humidi ...
from aerial parts, such as
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
, stems and
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
s. 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 Guttation is the exudation of drops of xylem sap on the tips or edges of leaves of some vascular plants, such as grasses, and a number of fungi, which are not plants but were previously categorized as such and studied as part of botany. Guttation ( ...
. Leaf surfaces are dotted with pores called
stoma In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
ta (singular "stoma"), and in most plants they are more numerous on the undersides of the foliage. The stomata are bordered by
guard cell Guard cells are specialized plant cells in the epidermis of leaves, stems and other organs that are used to control gas exchange. They are produced in pairs with a gap between them that forms a stomatal pore. The stomatal pores are largest when ...
s and their stomatal accessory cells (together known as stomatal complex) that open and close the pore. 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 Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
gas from the air for
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
. Transpiration also cools plants, changes
osmotic pressure Osmotic pressure is the minimum pressure which needs to be applied to a solution to prevent the inward flow of its pure solvent across a semipermeable membrane. It is also defined as the measure of the tendency of a solution to take in a pure ...
of cells, and enables
mass flow Mass flow, also known as mass transfer and bulk flow, is the movement of fluids down a pressure or temperature gradient,Moyes & Schulte (2008). Principles of Animal Physiology. Pearson Benjamin Cummings. San Francisco, California particularly in ...
of
mineral nutrients In the context of nutrition, a mineral is a chemical element required as an essential nutrient by organisms to perform functions necessary for life. However, the four major structural elements in the human body by weight (oxygen, hydrogen, carbon, ...
and water from roots to
shoot In botany, a plant shoot consists of any plant stem together with its appendages, leaves and lateral buds, flowering stems, and flower buds. The new growth from seed germination that grows upward is a shoot where leaves will develop. In the spri ...
s. 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. Mass flow of liquid water from the roots to the leaves is driven in part by
capillary action Capillary action (sometimes called capillarity, capillary motion, capillary rise, capillary effect, or wicking) is the process of a liquid flowing in a narrow space without the assistance of, or even in opposition to, any external forces li ...
, but primarily driven by
water potential Water potential is the potential energy of water per unit volume relative to pure water in reference conditions. Water potential quantifies the tendency of water to move from one area to another due to osmosis, gravity, mechanical pressure and mat ...
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 decreases 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. 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. In taller plants and trees, the force of gravity pulling the water inside can only be overcome by the decrease in
hydrostatic pressure Fluid statics or hydrostatics is the branch of fluid mechanics that studies the condition of the equilibrium of a floating body and submerged body "fluids at hydrostatic equilibrium and the pressure in a fluid, or exerted by a fluid, on an imme ...
in the upper parts of the plants due to the
diffusion Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemical p ...
of water out of stomata into the
atmosphere An atmosphere () is a layer of gas or layers of gases that envelop a planet, and is held in place by the gravity of the planetary body. A planet retains an atmosphere when the gravity is great and the temperature of the atmosphere is low. A s ...
. Water is absorbed by the roots by
osmosis Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of low water potential (region of ...
, and any dissolved mineral nutrients travel with it through the
xylem Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
. The
cohesion-tension theory Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
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.


Etymology

We can see the history of the word transpiration when we break it down into trans, a Latin noun that means "across," and spiration, which comes from the Latin verb spīrāre, meaning "to breathe." The motion suffix adds the meaning "the act of," so we can see transpiration is, literally, "the ACT of breathing across," which clearly identifies vapor emission from plant leaves.


Capillary action

Capillary action Capillary action (sometimes called capillarity, capillary motion, capillary rise, capillary effect, or wicking) is the process of a liquid flowing in a narrow space without the assistance of, or even in opposition to, any external forces li ...
is the process of a
liquid A liquid is a nearly incompressible fluid that conforms to the shape of its container but retains a (nearly) constant volume independent of pressure. As such, it is one of the four fundamental states of matter (the others being solid, gas, a ...
flowing in narrow spaces without the assistance of, or even in opposition to, external forces like
gravity In physics, gravity () is a fundamental interaction which causes mutual attraction between all things with mass or energy. Gravity is, by far, the weakest of the four fundamental interactions, approximately 1038 times weaker than the stro ...
. The effect can be seen in the drawing up of liquids between the hairs of a paint-brush, in a thin tube, in porous materials such as paper and plaster, in some non-porous materials such as sand and liquefied
carbon fiber Carbon fiber-reinforced polymers (American English), carbon-fibre-reinforced polymers (Commonwealth English), carbon-fiber-reinforced plastics, carbon-fiber reinforced-thermoplastic (CFRP, CRP, CFRTP), also known as carbon fiber, carbon compo ...
, or in a
biological cell The cell is the basic structural and functional unit of life forms. Every cell consists of a cytoplasm enclosed within a membrane, and contains many biomolecules such as proteins, DNA and RNA, as well as many small molecules of nutrients and ...
. It occurs because of
intermolecular force An intermolecular force (IMF) (or secondary force) is the force that mediates interaction between molecules, including the electromagnetic forces of attraction or repulsion which act between atoms and other types of neighbouring particles, e.g. a ...
s between the liquid and surrounding solid surfaces. If the diameter of the tube is sufficiently small, then the combination of
surface tension Surface tension is the tendency of liquid surfaces at rest to shrink into the minimum surface area possible. Surface tension is what allows objects with a higher density than water such as razor blades and insects (e.g. water striders) to f ...
(which is caused by cohesion within the liquid) and adhesive forces between the liquid and container wall act to propel the liquid.


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 Humidity is the concentration of water vapor present in the air. Water vapor, the gaseous state of water, is generally invisible to the human eye. Humidity indicates the likelihood for precipitation, dew, or fog to be present. Humidity depe ...
,
temperature Temperature is a physical quantity that expresses quantitatively the perceptions of hotness and coldness. Temperature is measured with a thermometer. Thermometers are calibrated in various temperature scales that historically have relied o ...
, wind, and incident sunlight. Along with above-ground factors, soil temperature and moisture can influence stomatal opening, 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 The enthalpy of vaporization (symbol ), also known as the (latent) heat of vaporization or heat of evaporation, is the amount of energy (enthalpy) that must be added to a liquid substance to transform a quantity of that substance into a gas. T ...
of 2260 kJ per liter. File: Transpiration Temperature Graph.svg, The effect of temperature on the transpiration rate of plants. File: Transpiration WindVelocity Graph.svg, The effect of wind velocity on the transpiration rate of plants. File: Transpiration Humidity Graph.svg, The effect of humidity on the transpiration rate of plants. 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
crop A crop is a plant that can be grown and harvested extensively for profit or subsistence. When the plants of the same kind are cultivated at one place on a large scale, it is called a crop. Most crops are cultivated in agriculture or hydroponic ...
s tends to fall between 200 and 1000 (''i.e.'', crop plants transpire 200 to 1000 kg of water for every kg of dry
matter In classical physics and general chemistry, matter is any substance that has mass and takes up space by having volume. All everyday objects that can be touched are ultimately composed of atoms, which are made up of interacting subatomic partic ...
produced). Transpiration rates of plants can be measured by a number of techniques, including potometers,
lysimeter A lysimeter (from Greek λύσις (loosening) and the suffix ''-meter'') is a measuring device which can be used to measure the amount of actual evapotranspiration which is released by plants (usually crops or trees). By recording the amount of p ...
s, porometers,
photosynthesis system Photosynthesis systems are electronic scientific instruments designed for non-destructive measurement of photosynthetic rates in the field. Photosynthesis systems are commonly used in agronomic and environmental research, as well as studies of t ...
s and thermometric sap flow sensors. Isotope measurements indicate transpiration is the larger component of
evapotranspiration Evapotranspiration (ET) is the combined processes by which water moves from the earth’s surface into the atmosphere. It covers both water evaporation (movement of water to the air directly from soil, canopies, and water bodies) and transpi ...
. Recent evidence from a global study 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.
Desert A desert is a barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to denudation. About on ...
plants have specially adapted structures, such as thick
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
s, reduced leaf areas, sunken stomata and
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...
s to reduce transpiration and conserve water. Many
cacti A cactus (, or less commonly, cactus) is a member of the plant family Cactaceae, a family comprising about 127 genera with some 1750 known species of the order Caryophyllales. The word ''cactus'' derives, through Latin, from the Ancient Greek ...
conduct
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
in
succulent In botany, succulent plants, also known as succulents, are plants with parts that are thickened, fleshy, and engorged, usually to retain water in arid climates or soil conditions. The word ''succulent'' comes from the Latin word ''sucus'', meani ...
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 Crassulacean acid metabolism, also known as CAM photosynthesis, is a carbon fixation pathway that evolved in some plants as an adaptation to arid conditions that allows a plant to photosynthesize during the day, but only exchange gases at night. ...
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 Cavitation is a phenomenon in which the static pressure of a liquid reduces to below the liquid's vapour pressure, leading to the formation of small vapor-filled cavities in the liquid. When subjected to higher pressure, these cavities, cal ...
occurs. 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. 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. 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. Scientists have begun using
magnetic resonance imaging Magnetic resonance imaging (MRI) is a medical imaging technique used in radiology to form pictures of the anatomy and the physiological processes of the body. MRI scanners use strong magnetic fields, magnetic field gradients, and radio wave ...
(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.


Effects on the environment


Cooling


See also

*
Antitranspirant Antitranspirants are compounds applied to the leaves of plants to reduce transpiration. They are used on Christmas trees, on cut flowers, on newly transplanted shrubs, and in other applications to preserve and protect plants from drying out too qu ...
– a substance to prevent transpiration *
Canopy conductance Canopy conductance, commonly denoted g_c, is a dimensionless quantity characterizing radiation distribution in tree canopy. By definition, it is calculated as a ratio of daily water use to daily mean vapor pressure deficit (VPD). Canopy conductanc ...
*
Ecohydrology Ecohydrology (from Greek , ''oikos'', "house(hold)"; , ''hydōr'', "water"; and , '' -logia'') is an interdisciplinary scientific field studying the interactions between water and ecological systems. It is considered a sub discipline of hydrology, ...
*
Eddy covariance The eddy covariance (also known as eddy correlation and eddy flux) is a key atmospheric measurement technique to measure and calculate vertical turbulent fluxes within atmospheric boundary layers. The method analyses high-frequency wind and scala ...
flux (aka eddy correlation, eddy flux) *
Hydrology (agriculture) Agricultural hydrology is the study of water balance components intervening in farm water, agricultural water management, especially in irrigation and drainage. Water balance components The water balance components can be grouped into components ...
*
Latent heat flux Latent heat (also known as latent energy or heat of transformation) is energy released or absorbed, by a body or a thermodynamic system, during a constant-temperature process — usually a first-order phase transition. Latent heat can be underst ...
*
Perspiration Perspiration, also known as sweating, is the production of fluids secreted by the sweat glands in the skin of mammals. Two types of sweat glands can be found in humans: eccrine glands and apocrine glands. The eccrine sweat glands are distrib ...
*
Soil plant atmosphere continuum The soil-plant-atmosphere continuum (SPAC) is the pathway for water moving from soil through plants to the atmosphere. Continuum in the description highlights the continuous nature of water connection through the pathway. The low water potential of ...
*
Stomatal conductance Stomatal conductance, usually measured in mmol m−2 s−1 by a porometer, estimates the rate of gas exchange (i.e., carbon dioxide uptake) and transpiration (i.e., water loss as water vapor) through the leaf stomata as determined by the degree of ...
*
Transpiration stream In plants, the transpiration stream is the uninterrupted stream of water and solutes which is taken up by the roots and transported via the xylem to the leaves where it evaporates into the air/apoplast-interface of the substomatal cavity. It is d ...
*
Turgor pressure Turgor pressure is the force within the cell that pushes the plasma membrane against the cell wall. It is also called ''hydrostatic pressure'', and is defined as the pressure in a fluid measured at a certain point within itself when at equilibri ...
* Water Evaluation And Planning system (WEAP)


References


External links


USGS The Water Cycle: Evapotranspiration
{{Authority control Hydrology Plant physiology