Transpiration is the process of

water Water is an inorganic compound with the chemical formula . It is a transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance. It is the main constituent of Earth's hydrosphere and the fluids of all known liv ...

movement through a

plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...

and its

evaporation Evaporation is a type of vaporization that occurs on the Interface (chemistry), surface of a liquid as it changes into the gas phase. A high concentration of the evaporating substance in the surrounding gas significantly slows down evapora ...

from aerial parts, such as

leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...

, stems and

flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...

s. It is a passive process that requires no energy expense by the plant. Transpiration also cools plants, changes

osmotic pressure Osmotic pressure is the minimum pressure which needs to be applied to a Solution (chemistry), 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 soluti ...

of cells, and enables mass flow of

mineral nutrients In the context of nutrition, a mineral is a chemical element. Some "minerals" are essential for life, but most are not. ''Minerals'' are one of the four groups of essential nutrients; the others are vitamins, essential fatty acids, and essenti ...

. When water uptake by the roots is less than the water lost to the

atmosphere An atmosphere () is a layer of gases that envelop an astronomical object, held in place by the gravity of the object. A planet retains an atmosphere when the gravity is great and the temperature of the atmosphere is low. A stellar atmosph ...

by evaporation, plants close small pores called stomata to decrease water loss, which slows down nutrient uptake and decreases CO₂ absorption from the atmosphere limiting metabolic processes,

photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...

, and growth.

Water and nutrient uptake

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 internal liquid out of the tips or edges of leaves of some vascular plants, and also a number of fungi. Ancient Latin gutta means "a drop of fluid", whence modern botany formed the word guttation to designa ...

. Water with any dissolved mineral nutrients is absorbed into the roots by

osmosis Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively permeable membrane, selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of ...

, which travels through the

xylem Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...

by way of water molecule

adhesion Adhesion is the tendency of dissimilar particles or interface (matter), surfaces to cling to one another. (Cohesion (chemistry), Cohesion refers to the tendency of similar or identical particles and surfaces to cling to one another.) The ...

and cohesion to the foliage and out small pores called

stoma In botany, a stoma (: stomata, from Greek language, Greek ''στόμα'', "mouth"), also called a stomate (: stomates), is a pore found in the Epidermis (botany), epidermis of leaves, stems, and other organs, that controls the rate of gas exc ...

ta (singular "stoma"). The stomata are bordered by

guard cell Guard cells are specialized cells in the epidermis of leaves, stems and other organs of land plants 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 lar ...

s and their stomatal accessory cells (together known as stomatal complex) that open and close the pore. The

cohesion-tension theory Xylem is one of the two types of transport tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts of the plants such ...

explains how leaves pull water through the xylem. Water molecules stick together or exhibit cohesion. As a water molecule evaporates from the leaf's surface it pulls on the adjacent water molecule, creating a continuous water flow through the plant. 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 external forces like Gravitation, gravity. The effe ...

, 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 that 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, thus exerting tension in the cells' water. 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. In taller plants and trees, the force of gravity pulling the water inside can only be overcome by the decrease in

hydrostatic pressure Hydrostatics is the branch of fluid mechanics that studies fluids at hydrostatic equilibrium and "the pressure in a fluid or exerted by a fluid on an immersed body". The word "hydrostatics" is sometimes used to refer specifically to water and o ...

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

Etymology

The word transpiration comes from the words trans, a Latin preposition that means "across," and spiration, which comes from the Latin verb spīrāre, meaning "to breathe." The motion suffix adds the meaning "the act of," creating the meaning, "the ACT of breathing across."

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 external forces like Gravitation, gravity. The effe ...

is the process of a

liquid Liquid is a state of matter with a definite volume but no fixed shape. Liquids adapt to the shape of their container and are nearly incompressible, maintaining their volume even under pressure. The density of a liquid is usually close to th ...

flowing in narrow spaces without the assistance of, or even in opposition to, external forces like

gravity In physics, gravity (), also known as gravitation or a gravitational interaction, is a fundamental interaction, a mutual attraction between all massive particles. On Earth, gravity takes a slightly different meaning: the observed force b ...

. 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 comp ...

, or in a

biological cell The cell is the basic structural and functional unit of all forms of life. Every cell consists of cytoplasm enclosed within a membrane; many cells contain organelles, each with a specific function. The term comes from the Latin word meaning ...

. It occurs because of

intermolecular force An intermolecular force (IMF; also 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. ...

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 (physics), tension is what allows objects with a higher density than water such as razor blades and insects (e.g. Ge ...

(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 (meteorology), precipitation, dew, or fog t ...

temperature Temperature is a physical quantity that quantitatively expresses the attribute of hotness or coldness. Temperature is measurement, measured with a thermometer. It reflects the average kinetic energy of the vibrating and colliding atoms making ...

, 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. Factors that effect root absorption of water include: moisture content of the soil, excessive soil fertility or salt content, poorly developed root systems, and those impacted by pathogenic bacteria and fungi such as

pythium ''Pythium'' is a genus of parasitic oomycetes. They were formerly classified as fungi. Most species are plant parasites, but '' Pythium insidiosum'' is an important pathogen of animals, causing pythiosis. The feet of the fungus gnat are fre ...

or rhizoctonia. 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 of water each day, and a large oak tree can transpire 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. In other words, a crop is a plant or plant product that is grown for a specific purpose such as food, Fiber, fibre, or fuel. When plants of the same spe ...

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 pa ...

produced). Transpiration rates of plants can be measured by a number of techniques, including potometers, lysimeters, 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 ...

s and thermometric sap flow sensors. Isotope measurements indicate transpiration is the larger component of

evapotranspiration Evapotranspiration (ET) refers to the combined processes which move water from the Earth's surface (open water and ice surfaces, bare soil and vegetation) into the Atmosphere of Earth, atmosphere. It covers both water evaporation (movement of w ...

. 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 landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...

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 ...

s to reduce transpiration and conserve water. Many

cacti A cactus (: cacti, cactuses, or less commonly, cactus) is a member of the plant family Cactaceae (), a family of the order Caryophyllales comprising about 127 genera with some 1,750 known species. The word ''cactus'' derives, through Latin, ...

conduct

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 in fluid mechanics and engineering normally is the phenomenon in which the static pressure of a liquid reduces to below the liquid's vapor pressure, leading to the formation of small vapor-filled cavities in the liquid. When sub ...

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 generate pictures of the anatomy and the physiological processes inside the body. MRI scanners use strong magnetic fields, magnetic field gradients, and ...

(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

Transpiration cools plants, as the evaporating water carries away heat energy due to its large

latent heat of vaporization In thermodynamics, 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 sub ...

of 2260 kJ per liter.

References

External links

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