Plant physiology
Encyclopedia
Plant physiology is a subdiscipline of botany
Botany
Botany, plant science, or plant biology is a branch of biology that involves the scientific study of plant life. Traditionally, botany also included the study of fungi, algae and viruses...

 concerned with the functioning, or physiology
Physiology
Physiology is the science of the function of living systems. This includes how organisms, organ systems, organs, cells, and bio-molecules carry out the chemical or physical functions that exist in a living system. The highest honor awarded in physiology is the Nobel Prize in Physiology or...

, of plant
Plant
Plants are living organisms belonging to the kingdom Plantae. Precise definitions of the kingdom vary, but as the term is used here, plants include familiar organisms such as trees, flowers, herbs, bushes, grasses, vines, ferns, mosses, and green algae. The group is also called green plants or...

s. Closely related fields include plant morphology
Plant morphology
Plant morphology or phytomorphology is the study of the physical form and external structure of plants. This is usually considered distinct from plant anatomy, which is the study of the internal structure of plants, especially at the microscopic level...

 (structure of plants), plant ecology
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...

 (interactions with the environment), phytochemistry
Phytochemistry
Phytochemistry is in the strict sense of the word the study of phytochemicals. These are chemicals derived from plants. In a narrower sense the terms are often used to describe the large number of secondary metabolic compounds found in plants. Many of these are known to provide protection against...

 (biochemistry
Biochemistry
Biochemistry, sometimes called biological chemistry, is the study of chemical processes in living organisms, including, but not limited to, living matter. Biochemistry governs all living organisms and living processes...

 of plants), cell biology
Cell biology
Cell biology is a scientific discipline that studies cells – their physiological properties, their structure, the organelles they contain, interactions with their environment, their life cycle, division and death. This is done both on a microscopic and molecular level...

, and molecular biology
Molecular biology
Molecular biology is the branch of biology that deals with the molecular basis of biological activity. This field overlaps with other areas of biology and chemistry, particularly genetics and biochemistry...

.

Fundamental processes such as photosynthesis
Photosynthesis
Photosynthesis is a chemical process that converts carbon dioxide into organic compounds, especially sugars, using the energy from sunlight. Photosynthesis occurs in plants, algae, and many species of bacteria, but not in archaea. Photosynthetic organisms are called photoautotrophs, since they can...

, respiration
Cellular respiration
Cellular respiration is the set of the metabolic reactions and processes that take place in the cells of organisms to convert biochemical energy from nutrients into adenosine triphosphate , and then release waste products. The reactions involved in respiration are catabolic reactions that involve...

, plant nutrition
Plant nutrition
'Plant Nutrition is the study of the chemical elements that are necessary for growth. In 1972, E. Epstein defined 2 criteria for an element to be essential for plant growth:# in its absence the plant is unable to complete a normal life cycle or...

, plant hormone
Plant hormone
Plant hormones are chemicals that regulate plant growth, which, in the UK, are termed 'plant growth substances'. Plant hormones are signal molecules produced within the plant, and occur in extremely low concentrations. Hormones regulate cellular processes in targeted cells locally and, when moved...

 functions, tropism
Tropism
A tropism is a biological phenomenon, indicating growth or turning movement of a biological organism, usually a plant, in response to an environmental stimulus. In tropisms, this response is dependent on the direction of the stimulus...

s, nastic movements
Nastic movements
Nastic movements are non-directional responses to stimuli . The movement can be due to changes in turgor or changes in oma. Nastic movements differ banaan tropic movements in that the direction of tropic responses depends on the direction of the stimulus, whereas the direction of nastic movements...

, photoperiodism
Photoperiodism
Photoperiodism is the physiological reaction of organisms to the length of day or night. It occurs in plants and animals.Photoperiodism can also be defined as the developmental responses of plants to the relative lengths of the light and dark periods...

, photomorphogenesis
Photomorphogenesis
In developmental biology, photomorphogenesis is light-mediated development. The photomorphogenesis of plants is often studied by using tightly-frequency-controlled light sources to grow the plants.-Germination:...

, circadian rhythms, environmental stress physiology, seed germination
Germination
Germination is the process in which a plant or fungus emerges from a seed or spore, respectively, and begins growth. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However the growth of a sporeling from a spore, for example the...

, dormancy
Dormancy
Dormancy is a period in an organism's life cycle when growth, development, and physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be closely associated with environmental conditions...

 and stoma
Stoma
In botany, a stoma is a pore, found in the leaf and stem epidermis that is used forgas exchange. The pore is bordered by a pair of specialized parenchyma cells known as guard cells that are responsible for regulating the size of the opening...

ta function and transpiration
Transpiration
Transpiration is a process similar to evaporation. It is a part of the water cycle, and it is the loss of water vapor from parts of plants , especially in leaves but also in stems, flowers and roots. Leaf surfaces are dotted with openings which are collectively called stomata, and in most plants...

, both part of plant water relations, are studied by plant physiologists.

Scope

The field of plant physiology includes the study of all the internal activities of plants—those chemical and physical processes associated with life
Life
Life is a characteristic that distinguishes objects that have signaling and self-sustaining processes from those that do not, either because such functions have ceased , or else because they lack such functions and are classified as inanimate...

 as they occur in plants. This includes study at many levels of scale of size and time. At the smallest scale are molecular
Molecule
A molecule is an electrically neutral group of at least two atoms held together by covalent chemical bonds. Molecules are distinguished from ions by their electrical charge...

 interactions of photosynthesis
Photosynthesis
Photosynthesis is a chemical process that converts carbon dioxide into organic compounds, especially sugars, using the energy from sunlight. Photosynthesis occurs in plants, algae, and many species of bacteria, but not in archaea. Photosynthetic organisms are called photoautotrophs, since they can...

 and internal diffusion
Diffusion
Molecular diffusion, often called simply diffusion, is the thermal motion of all particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size of the particles...

 of water, minerals, and nutrients. At the largest scale are the processes of plant development
Developmental biology
Developmental biology is the study of the process by which organisms grow and develop. Modern developmental biology studies the genetic control of cell growth, differentiation and "morphogenesis", which is the process that gives rise to tissues, organs and anatomy.- Related fields of study...

, season
Season
A season is a division of the year, marked by changes in weather, ecology, and hours of daylight.Seasons result from the yearly revolution of the Earth around the Sun and the tilt of the Earth's axis relative to the plane of revolution...

ality, dormancy
Dormancy
Dormancy is a period in an organism's life cycle when growth, development, and physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be closely associated with environmental conditions...

, and reproductive control. Major subdisciplines of plant physiology include phytochemistry
Phytochemistry
Phytochemistry is in the strict sense of the word the study of phytochemicals. These are chemicals derived from plants. In a narrower sense the terms are often used to describe the large number of secondary metabolic compounds found in plants. Many of these are known to provide protection against...

 (the study of the biochemistry
Biochemistry
Biochemistry, sometimes called biological chemistry, is the study of chemical processes in living organisms, including, but not limited to, living matter. Biochemistry governs all living organisms and living processes...

 of plants) and phytopathology
Phytopathology
Plant pathology is the scientific study of plant diseases caused by pathogens and environmental conditions . Organisms that cause infectious disease include fungi, oomycetes, bacteria, viruses, viroids, virus-like organisms, phytoplasmas, protozoa, nematodes and parasitic plants...

 (the study of disease
Disease
A disease is an abnormal condition affecting the body of an organism. It is often construed to be a medical condition associated with specific symptoms and signs. It may be caused by external factors, such as infectious disease, or it may be caused by internal dysfunctions, such as autoimmune...

 in plants). The scope of plant physiology as a discipline may be divided into several major areas of research.

First, the study of phytochemistry
Phytochemistry
Phytochemistry is in the strict sense of the word the study of phytochemicals. These are chemicals derived from plants. In a narrower sense the terms are often used to describe the large number of secondary metabolic compounds found in plants. Many of these are known to provide protection against...

 (plant chemistry) is included within the domain of plant physiology. To function and survive, plants produce a wide array of chemical compounds not found in other organisms. Photosynthesis
Photosynthesis
Photosynthesis is a chemical process that converts carbon dioxide into organic compounds, especially sugars, using the energy from sunlight. Photosynthesis occurs in plants, algae, and many species of bacteria, but not in archaea. Photosynthetic organisms are called photoautotrophs, since they can...

 requires a large array of pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...

s, enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...

s, and other compounds to function. Because they cannot move, plants must also defend themselves chemically from herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...

s, pathogen
Pathogen
A pathogen gignomai "I give birth to") or infectious agent — colloquially, a germ — is a microbe or microorganism such as a virus, bacterium, prion, or fungus that causes disease in its animal or plant host...

s and competition from other plants. They do this by producing toxin
Toxin
A toxin is a poisonous substance produced within living cells or organisms; man-made substances created by artificial processes are thus excluded...

s and foul-tasting or smelling chemicals. Other compounds defend plants against disease, permit survival during drought, and prepare plants for dormancy. While other compounds are used to attract pollinator
Pollinator
A pollinator is the biotic agent that moves pollen from the male anthers of a flower to the female stigma of a flower to accomplish fertilization or syngamy of the female gamete in the ovule of the flower by the male gamete from the pollen grain...

s or herbivores to spread ripe seeds.

Secondly, plant physiology includes the study of biological and chemical processes of individual plant cells
Cell (biology)
The cell is the basic structural and functional unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often called the building block of life. The Alberts text discusses how the "cellular building blocks" move to shape developing embryos....

. Plant cells have a number of features that distinguish them from cells of animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...

s, and which lead to major differences in the way that plant life behaves and responds differently from animal life. For example, plant cells have a cell wall
Cell wall
The cell wall is the tough, usually flexible but sometimes fairly rigid layer that surrounds some types of cells. It is located outside the cell membrane and provides these cells with structural support and protection, and also acts as a filtering mechanism. A major function of the cell wall is to...

 which restricts the shape of plant cells and thereby limits the flexibility and mobility of plants. Plant cells also contain chlorophyll
Chlorophyll
Chlorophyll is a green pigment found in almost all plants, algae, and cyanobacteria. Its name is derived from the Greek words χλωρος, chloros and φύλλον, phyllon . Chlorophyll is an extremely important biomolecule, critical in photosynthesis, which allows plants to obtain energy from light...

, a chemical compound that interacts with light
Light
Light or visible light is electromagnetic radiation that is visible to the human eye, and is responsible for the sense of sight. Visible light has wavelength in a range from about 380 nanometres to about 740 nm, with a frequency range of about 405 THz to 790 THz...

 in a way that enables plants to manufacture their own nutrients rather than consuming other living things as animals do.

Thirdly, plant physiology deals with interactions between cells, tissues
Tissue (biology)
Tissue is a cellular organizational level intermediate between cells and a complete organism. A tissue is an ensemble of cells, not necessarily identical, but from the same origin, that together carry out a specific function. These are called tissues because of their identical functioning...

, and organs within a plant. Different cells and tissues are physically and chemically specialized to perform different functions. Root
Root
In vascular plants, the root is the organ of a plant that typically lies below the surface of the soil. This is not always the case, however, since a root can also be aerial or aerating . Furthermore, a stem normally occurring below ground is not exceptional either...

s and rhizoid
Rhizoid
Rhizoids are thread-like growths from the base or bottom of a plant, found mainly in lower groups such as algae, fungi, bryophytes and pteridophytes, that function like roots of higher plants ....

s function to anchor the plant and acquire minerals in the soil. Leaves
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

 catch light in order to manufacture nutrients. For both of these organs to remain living, minerals that the roots acquire must be transported to the leaves, and the nutrients manufactured in the leaves must be transported to the roots. Plants have developed a number of ways to achieve this transport, such as vascular tissue
Vascular tissue
Vascular tissue is a complex conducting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue are the xylem and phloem. These two tissues transport fluid and nutrients internally. There are also two meristems associated with vascular tissue:...

, and the functioning of the various modes of transport is studied by plant physiologists.

Fourthly, plant physiologists study the ways that plants control or regulate internal functions. Like animals, plants produce chemicals called hormones
Plant hormone
Plant hormones are chemicals that regulate plant growth, which, in the UK, are termed 'plant growth substances'. Plant hormones are signal molecules produced within the plant, and occur in extremely low concentrations. Hormones regulate cellular processes in targeted cells locally and, when moved...

 which are produced in one part of the plant to signal cells in another part of the plant to respond. Many flowering plant
Flowering plant
The flowering plants , also known as Angiospermae or Magnoliophyta, are the most diverse group of land plants. Angiosperms are seed-producing plants like the gymnosperms and can be distinguished from the gymnosperms by a series of synapomorphies...

s bloom at the appropriate time because of light-sensitive compounds that respond to the length of the night, a phenomenon known as photoperiodism
Photoperiodism
Photoperiodism is the physiological reaction of organisms to the length of day or night. It occurs in plants and animals.Photoperiodism can also be defined as the developmental responses of plants to the relative lengths of the light and dark periods...

. The ripening
Ripening
Ripening is a process in fruits that causes them to become more palatable. In general, a fruit becomes sweeter, less green, and softer as it ripens. Even though the acidity of fruit increases as it ripens, the higher acidity level does not make the fruit seem tarter, which can lead to the...

 of fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...

 and loss of leaves in the winter are controlled in part by the production of the gas ethylene
Ethylene
Ethylene is a gaseous organic compound with the formula . It is the simplest alkene . Because it contains a carbon-carbon double bond, ethylene is classified as an unsaturated hydrocarbon. Ethylene is widely used in industry and is also a plant hormone...

 by the plant.

Finally, plant physiology includes the study of how plants respond to conditions and variation in the environment, a field known as environmental physiology. Stress from water loss, changes in air chemistry, or crowding by other plants can lead to changes in the way a plant functions. These changes may be affected by genetic, chemical, and physical factors.

Biochemistry of plants

The list of simple elements
Chemical element
A chemical element is a pure chemical substance consisting of one type of atom distinguished by its atomic number, which is the number of protons in its nucleus. Familiar examples of elements include carbon, oxygen, aluminum, iron, copper, gold, mercury, and lead.As of November 2011, 118 elements...

 of which plants are primarily constructed—carbon
Carbon
Carbon is the chemical element with symbol C and atomic number 6. As a member of group 14 on the periodic table, it is nonmetallic and tetravalent—making four electrons available to form covalent chemical bonds...

, oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...

, hydrogen
Hydrogen
Hydrogen is the chemical element with atomic number 1. It is represented by the symbol H. With an average atomic weight of , hydrogen is the lightest and most abundant chemical element, constituting roughly 75% of the Universe's chemical elemental mass. Stars in the main sequence are mainly...

, calcium
Calcium
Calcium is the chemical element with the symbol Ca and atomic number 20. It has an atomic mass of 40.078 amu. Calcium is a soft gray alkaline earth metal, and is the fifth-most-abundant element by mass in the Earth's crust...

, phosphorus
Phosphorus
Phosphorus is the chemical element that has the symbol P and atomic number 15. A multivalent nonmetal of the nitrogen group, phosphorus as a mineral is almost always present in its maximally oxidized state, as inorganic phosphate rocks...

, etc.—is not different from similar lists for animals, fungi, or even bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...

. The fundamental atomic components of plants are the same as for all life; only the details of the way in which they are assembled differs.

Despite this underlying similarity, plants produce a vast array of chemical compounds with unusual properties which they use to cope with their environment. Pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...

s are used by plants to absorb or detect light, and are extracted by humans for use in dye
Dye
A dye is a colored substance that has an affinity to the substrate to which it is being applied. The dye is generally applied in an aqueous solution, and requires a mordant to improve the fastness of the dye on the fiber....

s. Other plant products may be used for the manufacture of commercially important rubber
Rubber
Natural rubber, also called India rubber or caoutchouc, is an elastomer that was originally derived from latex, a milky colloid produced by some plants. The plants would be ‘tapped’, that is, an incision made into the bark of the tree and the sticky, milk colored latex sap collected and refined...

 or biofuel
Biofuel
Biofuel is a type of fuel whose energy is derived from biological carbon fixation. Biofuels include fuels derived from biomass conversion, as well as solid biomass, liquid fuels and various biogases...

. Perhaps the most celebrated compounds from plants are those with pharmacological activity, such as salicylic acid
Salicylic acid
Salicylic acid is a monohydroxybenzoic acid, a type of phenolic acid and a beta hydroxy acid. This colorless crystalline organic acid is widely used in organic synthesis and functions as a plant hormone. It is derived from the metabolism of salicin...

 (aspirin), morphine
Morphine
Morphine is a potent opiate analgesic medication and is considered to be the prototypical opioid. It was first isolated in 1804 by Friedrich Sertürner, first distributed by same in 1817, and first commercially sold by Merck in 1827, which at the time was a single small chemists' shop. It was more...

, and digitalis
Digitalis
Digitalis is a genus of about 20 species of herbaceous perennials, shrubs, and biennials that are commonly called foxgloves. This genus was traditionally placed in the figwort family Scrophulariaceae, but recent reviews of phylogenetic research have placed it in the much enlarged family...

. Drug companies spend billions of dollars each year researching plant compounds for potential medicinal benefits.

Constituent elements

Plants require some nutrient
Nutrient
A nutrient is a chemical that an organism needs to live and grow or a substance used in an organism's metabolism which must be taken in from its environment. They are used to build and repair tissues, regulate body processes and are converted to and used as energy...

s, such as carbon
Carbon
Carbon is the chemical element with symbol C and atomic number 6. As a member of group 14 on the periodic table, it is nonmetallic and tetravalent—making four electrons available to form covalent chemical bonds...

 and nitrogen
Nitrogen
Nitrogen is a chemical element that has the symbol N, atomic number of 7 and atomic mass 14.00674 u. Elemental nitrogen is a colorless, odorless, tasteless, and mostly inert diatomic gas at standard conditions, constituting 78.08% by volume of Earth's atmosphere...

, in large quantities to survive. Such nutrients are termed macronutrients, where the prefix macro- (large) refers to the quantity needed, not the size of the nutrient particles themselves. Other nutrients, called micronutrient
Micronutrient
Micronutrients are nutrients required by humans and other living things throughout life in small quantities to orchestrate a whole range of physiological functions, but which the organism itself cannot produce. For people, they include dietary trace minerals in amounts generally less than 100...

s, are required only in trace amounts for plants to remain healthy. Such micronutrients are usually absorbed as ion
Ion
An ion is an atom or molecule in which the total number of electrons is not equal to the total number of protons, giving it a net positive or negative electrical charge. The name was given by physicist Michael Faraday for the substances that allow a current to pass between electrodes in a...

s dissolved in water taken from the soil, though carnivorous plant
Carnivorous plant
Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans, typically insects and other arthropods. Carnivorous plants appear adapted to grow in places where the soil is thin or poor in nutrients, especially nitrogen, such as acidic...

s acquire some of their micronutrients from captured prey.

The following tables list element
Chemical element
A chemical element is a pure chemical substance consisting of one type of atom distinguished by its atomic number, which is the number of protons in its nucleus. Familiar examples of elements include carbon, oxygen, aluminum, iron, copper, gold, mercury, and lead.As of November 2011, 118 elements...

 nutrients essential to plants. Uses within plants are generalized.
Macronutrients. (Necessary in large quantities)
Element Form of uptake Notes
Nitrogen
Nitrogen
Nitrogen is a chemical element that has the symbol N, atomic number of 7 and atomic mass 14.00674 u. Elemental nitrogen is a colorless, odorless, tasteless, and mostly inert diatomic gas at standard conditions, constituting 78.08% by volume of Earth's atmosphere...

 
NO3, NH4+ Nucleic acids, proteins, hormones, etc.
Oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...

 
O2 H2O Cellulose
Cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to over ten thousand β linked D-glucose units....

, starch
Starch
Starch or amylum is a carbohydrate consisting of a large number of glucose units joined together by glycosidic bonds. This polysaccharide is produced by all green plants as an energy store...

, other organic compounds
Carbon
Carbon
Carbon is the chemical element with symbol C and atomic number 6. As a member of group 14 on the periodic table, it is nonmetallic and tetravalent—making four electrons available to form covalent chemical bonds...

 
CO2 Cellulose, starch, other organic compounds
Hydrogen
Hydrogen
Hydrogen is the chemical element with atomic number 1. It is represented by the symbol H. With an average atomic weight of , hydrogen is the lightest and most abundant chemical element, constituting roughly 75% of the Universe's chemical elemental mass. Stars in the main sequence are mainly...

 
H2O Cellulose, starch, other organic compounds
Potassium
Potassium
Potassium is the chemical element with the symbol K and atomic number 19. Elemental potassium is a soft silvery-white alkali metal that oxidizes rapidly in air and is very reactive with water, generating sufficient heat to ignite the hydrogen emitted in the reaction.Potassium and sodium are...

 
K+ Cofactor in protein synthesis, water balance, etc.
Calcium
Calcium
Calcium is the chemical element with the symbol Ca and atomic number 20. It has an atomic mass of 40.078 amu. Calcium is a soft gray alkaline earth metal, and is the fifth-most-abundant element by mass in the Earth's crust...

 
Ca2+ Membrane synthesis and stabilization
Magnesium
Magnesium
Magnesium is a chemical element with the symbol Mg, atomic number 12, and common oxidation number +2. It is an alkaline earth metal and the eighth most abundant element in the Earth's crust and ninth in the known universe as a whole...

 
Mg2+ Element essential for chlorophyll
Phosphorus
Phosphorus
Phosphorus is the chemical element that has the symbol P and atomic number 15. A multivalent nonmetal of the nitrogen group, phosphorus as a mineral is almost always present in its maximally oxidized state, as inorganic phosphate rocks...

 
H2PO4 Nucleic acids, phospholipids, ATP
Sulfur
Sulfur
Sulfur or sulphur is the chemical element with atomic number 16. In the periodic table it is represented by the symbol S. It is an abundant, multivalent non-metal. Under normal conditions, sulfur atoms form cyclic octatomic molecules with chemical formula S8. Elemental sulfur is a bright yellow...

 
SO42– Constituent of proteins and coenzymes

Micronutrients. (Necessary in small quantities)
Element Form of uptake Notes
Chlorine
Chlorine
Chlorine is the chemical element with atomic number 17 and symbol Cl. It is the second lightest halogen, found in the periodic table in group 17. The element forms diatomic molecules under standard conditions, called dichlorine...

 
Cl- Photosystem II and stomata function
Iron
Iron
Iron is a chemical element with the symbol Fe and atomic number 26. It is a metal in the first transition series. It is the most common element forming the planet Earth as a whole, forming much of Earth's outer and inner core. It is the fourth most common element in the Earth's crust...

 
Fe2+, Fe3+ Chorophyll formation
Boron
Boron
Boron is the chemical element with atomic number 5 and the chemical symbol B. Boron is a metalloid. Because boron is not produced by stellar nucleosynthesis, it is a low-abundance element in both the solar system and the Earth's crust. However, boron is concentrated on Earth by the...

 
HBO3 Crosslinking pectin
Manganese
Manganese
Manganese is a chemical element, designated by the symbol Mn. It has the atomic number 25. It is found as a free element in nature , and in many minerals...

 
Mn2+ Activity of some enzymes
Zinc
Zinc
Zinc , or spelter , is a metallic chemical element; it has the symbol Zn and atomic number 30. It is the first element in group 12 of the periodic table. Zinc is, in some respects, chemically similar to magnesium, because its ion is of similar size and its only common oxidation state is +2...

 
Zn2+ Involved in the synthesis of enzymes and chlorophyll
Copper
Copper
Copper is a chemical element with the symbol Cu and atomic number 29. It is a ductile metal with very high thermal and electrical conductivity. Pure copper is soft and malleable; an exposed surface has a reddish-orange tarnish...

 
Cu+ Enzymes for lignin synthesis
Molybdenum
Molybdenum
Molybdenum , is a Group 6 chemical element with the symbol Mo and atomic number 42. The name is from Neo-Latin Molybdaenum, from Ancient Greek , meaning lead, itself proposed as a loanword from Anatolian Luvian and Lydian languages, since its ores were confused with lead ores...

 
MoO42- Nitrogen fixation, reduction of nitrates
Nickel
Nickel
Nickel is a chemical element with the chemical symbol Ni and atomic number 28. It is a silvery-white lustrous metal with a slight golden tinge. Nickel belongs to the transition metals and is hard and ductile...

 
Ni2+ Enzymatic cofactor in the metabolism of nitrogen compounds

Pigments

Among the most important molecules for plant function are the pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...

s. Plant pigments include a variety of different kinds of molecules, including porphyrin
Porphyrin
Porphyrins are a group of organic compounds, many naturally occurring. One of the best-known porphyrins is heme, the pigment in red blood cells; heme is a cofactor of the protein hemoglobin. Porphyrins are heterocyclic macrocycles composed of four modified pyrrole subunits interconnected at...

s, carotenoid
Carotenoid
Carotenoids are tetraterpenoid organic pigments that are naturally occurring in the chloroplasts and chromoplasts of plants and some other photosynthetic organisms like algae, some bacteria, and some types of fungus. Carotenoids can be synthesized fats and other basic organic metabolic building...

s, and anthocyanin
Anthocyanin
Anthocyanins are water-soluble vacuolar pigments that may appear red, purple, or blue according to pH...

s. All biological pigment
Biological pigment
Biological pigments, also known simply as pigments or biochromes are substances produced by living organisms that have a color resulting from selective color absorption. Biological pigments include plant pigments and flower pigments...

s selectively absorb certain wavelength
Wavelength
In physics, the wavelength of a sinusoidal wave is the spatial period of the wave—the distance over which the wave's shape repeats.It is usually determined by considering the distance between consecutive corresponding points of the same phase, such as crests, troughs, or zero crossings, and is a...

s of light
Light
Light or visible light is electromagnetic radiation that is visible to the human eye, and is responsible for the sense of sight. Visible light has wavelength in a range from about 380 nanometres to about 740 nm, with a frequency range of about 405 THz to 790 THz...

 while reflect
Reflection (physics)
Reflection is the change in direction of a wavefront at an interface between two differentmedia so that the wavefront returns into the medium from which it originated. Common examples include the reflection of light, sound and water waves...

ing others. The light that is absorbed may be used by the plant to power chemical reaction
Chemical reaction
A chemical reaction is a process that leads to the transformation of one set of chemical substances to another. Chemical reactions can be either spontaneous, requiring no input of energy, or non-spontaneous, typically following the input of some type of energy, such as heat, light or electricity...

s, while the reflected wavelengths of light determine the color
Color
Color or colour is the visual perceptual property corresponding in humans to the categories called red, green, blue and others. Color derives from the spectrum of light interacting in the eye with the spectral sensitivities of the light receptors...

 the pigment appears to the eye.

Chlorophyll
Chlorophyll
Chlorophyll is a green pigment found in almost all plants, algae, and cyanobacteria. Its name is derived from the Greek words χλωρος, chloros and φύλλον, phyllon . Chlorophyll is an extremely important biomolecule, critical in photosynthesis, which allows plants to obtain energy from light...

 is the primary pigment in plants; it is a porphyrin
Porphyrin
Porphyrins are a group of organic compounds, many naturally occurring. One of the best-known porphyrins is heme, the pigment in red blood cells; heme is a cofactor of the protein hemoglobin. Porphyrins are heterocyclic macrocycles composed of four modified pyrrole subunits interconnected at...

 that absorbs red and blue wavelengths of light while reflecting green
Green
Green is a color, the perception of which is evoked by light having a spectrum dominated by energy with a wavelength of roughly 520–570 nanometres. In the subtractive color system, it is not a primary color, but is created out of a mixture of yellow and blue, or yellow and cyan; it is considered...

. It is the presence and relative abundance of chlorophyll that gives plants their green color. All land plants and green algae possess two forms of this pigment: chlorophyll a and chlorophyll b. Kelp
Kelp
Kelps are large seaweeds belonging to the brown algae in the order Laminariales. There are about 30 different genera....

s, diatom
Diatom
Diatoms are a major group of algae, and are one of the most common types of phytoplankton. Most diatoms are unicellular, although they can exist as colonies in the shape of filaments or ribbons , fans , zigzags , or stellate colonies . Diatoms are producers within the food chain...

s, and other photosynthetic heterokont
Heterokont
The heterokonts or stramenopiles are a major line of eukaryotes currently containing more than 100,000 known species. Most are algae, ranging from the giant multicellular kelp to the unicellular diatoms, which are a primary component of plankton...

s contain chlorophyll c instead of b, while red algae
Red algae
The red algae are one of the oldest groups of eukaryotic algae, and also one of the largest, with about 5,000–6,000 species  of mostly multicellular, marine algae, including many notable seaweeds...

 possess only chlorophyll a. All chlorophylls serve as the primary means plants use to intercept light to fuel photosynthesis
Photosynthesis
Photosynthesis is a chemical process that converts carbon dioxide into organic compounds, especially sugars, using the energy from sunlight. Photosynthesis occurs in plants, algae, and many species of bacteria, but not in archaea. Photosynthetic organisms are called photoautotrophs, since they can...

.

Carotenoid
Carotenoid
Carotenoids are tetraterpenoid organic pigments that are naturally occurring in the chloroplasts and chromoplasts of plants and some other photosynthetic organisms like algae, some bacteria, and some types of fungus. Carotenoids can be synthesized fats and other basic organic metabolic building...

s are red, orange, or yellow tetraterpenoid
Tetraterpenoid
Tetraterpenoids are terpenoids of 8 isoprene units - hence 40 carbon atoms in the skeleton.Notable subgroups are*carotenoidsOthers*trianthenol 1*methyl tortuoate C from coral...

s. They function as accessory pigments in plants, helping to fuel photosynthesis
Photosynthesis
Photosynthesis is a chemical process that converts carbon dioxide into organic compounds, especially sugars, using the energy from sunlight. Photosynthesis occurs in plants, algae, and many species of bacteria, but not in archaea. Photosynthetic organisms are called photoautotrophs, since they can...

 by gathering wavelengths of light not readily absorbed by chlorophyll. The most familiar carotenoids are carotene
Carotene
The term carotene is used for several related unsaturated hydrocarbon substances having the formula C40Hx, which are synthesized by plants but cannot be made by animals. Carotene is an orange photosynthetic pigment important for photosynthesis. Carotenes are all coloured to the human eye...

 (an orange pigment found in carrot
Carrot
The carrot is a root vegetable, usually orange in colour, though purple, red, white, and yellow varieties exist. It has a crisp texture when fresh...

s), lutein
Lutein
Lutein is a xanthophyll and one of 600 known naturally occurring carotenoids. Lutein is synthesized only by plants and like other xanthophylls is found in high quantities in green leafy vegetables such as spinach and kale...

 (a yellow pigment found in fruits and vegetables), and lycopene
Lycopene
Lycopene is a bright red carotene and carotenoid pigment and phytochemical found in tomatoes and other red fruits and vegetables, such as red carrots, watermelons and papayas...

 (the red pigment responsible for the color of tomato
Tomato
The word "tomato" may refer to the plant or the edible, typically red, fruit which it bears. Originating in South America, the tomato was spread around the world following the Spanish colonization of the Americas, and its many varieties are now widely grown, often in greenhouses in cooler...

es). Carotenoids have been shown to act as antioxidant
Antioxidant
An antioxidant is a molecule capable of inhibiting the oxidation of other molecules. Oxidation is a chemical reaction that transfers electrons or hydrogen from a substance to an oxidizing agent. Oxidation reactions can produce free radicals. In turn, these radicals can start chain reactions. When...

s and to promote healthy eyesight in humans.

Anthocyanin
Anthocyanin
Anthocyanins are water-soluble vacuolar pigments that may appear red, purple, or blue according to pH...

s (literally "flower blue") are water-soluble
Solubility
Solubility is the property of a solid, liquid, or gaseous chemical substance called solute to dissolve in a solid, liquid, or gaseous solvent to form a homogeneous solution of the solute in the solvent. The solubility of a substance fundamentally depends on the used solvent as well as on...

 flavonoid
Flavonoid
Flavonoids , are a class of plant secondary metabolites....

 pigments that appear red to blue, according to pH
PH
In chemistry, pH is a measure of the acidity or basicity of an aqueous solution. Pure water is said to be neutral, with a pH close to 7.0 at . Solutions with a pH less than 7 are said to be acidic and solutions with a pH greater than 7 are basic or alkaline...

. They occur in all tissues
Tissue (biology)
Tissue is a cellular organizational level intermediate between cells and a complete organism. A tissue is an ensemble of cells, not necessarily identical, but from the same origin, that together carry out a specific function. These are called tissues because of their identical functioning...

 of higher plants, providing color in leaves
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

, stems
Plant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...

, root
Root
In vascular plants, the root is the organ of a plant that typically lies below the surface of the soil. This is not always the case, however, since a root can also be aerial or aerating . Furthermore, a stem normally occurring below ground is not exceptional either...

s, flower
Flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...

s, and fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...

s, though not always in sufficient quantities to be noticeable. Anthocyanins are most visible in the petal
Petal
Petals are modified leaves that surround the reproductive parts of flowers. They often are brightly colored or unusually shaped to attract pollinators. Together, all of the petals of a flower are called a corolla. Petals are usually accompanied by another set of special leaves called sepals lying...

s of flowers, where they may make up as much as 30% of the dry weight of the tissue. They are also responsible for the purple color seen on the underside of tropical shade plants such as Tradescantia zebrina
Tradescantia zebrina
Tradescantia zebrina, formerly known as Zebrina pendula, is a species of spiderwort more commonly known as an inch plant or wandering jew, native to the Gulf Coast region of eastern Mexico. The common name is shared with closely related varieties T. fluminensis and T...

. In these plants, the anthocyanin catches light that has passed through the leaf and reflects it back towards regions bearing chlorophyll, in order to maximize the use of available light

Betalain
Betalain
Betalains are a class of red and yellow indole-derived pigments found in plants of the Caryophyllales, where they replace anthocyanin pigments. Betalains also occur in some higher order fungi. They are most often noticeable in the petals of flowers, but may color the fruits, leaves, stems, and...

s are red or yellow pigments. Like anthocyanins they are water-soluble, but unlike anthocyanins they are indole
Indole
Indole is an aromatic heterocyclic organic compound. It has a bicyclic structure, consisting of a six-membered benzene ring fused to a five-membered nitrogen-containing pyrrole ring. Indole is a popular component of fragrances and the precursor to many pharmaceuticals. Compounds that contain an...

-derived compounds synthesized from tyrosine
Tyrosine
Tyrosine or 4-hydroxyphenylalanine, is one of the 22 amino acids that are used by cells to synthesize proteins. Its codons are UAC and UAU. It is a non-essential amino acid with a polar side group...

. This class of pigments is found only in the Caryophyllales
Caryophyllales
Caryophyllales is an order of flowering plants that includes the cacti, carnations, amaranths, ice plants, and many carnivorous plants. Many members are succulent, having fleshy stems or leaves.-Description:...

 (including cactus
Cactus
A cactus is a member of the plant family Cactaceae. Their distinctive appearance is a result of adaptations to conserve water in dry and/or hot environments. In most species, the stem has evolved to become photosynthetic and succulent, while the leaves have evolved into spines...

 and amaranth
Amaranth
Amaranthus, collectively known as amaranth, is a cosmopolitan genus of herbs. Approximately 60 species are recognized, with inflorescences and foliage ranging from purple and red to gold...

), and never co-occur in plants with anthocyanins. Betalains are responsible for the deep red color of beet
Beet
The beet is a plant in the Chenopodiaceae family which is now included in Amaranthaceae family. It is best known in its numerous cultivated varieties, the most well known of which is the purple root vegetable known as the beetroot or garden beet...

s, and are used commercially as food-coloring agents. Plant physiologists are uncertain of the function that betalains have in plants which possess them, but there is some preliminary evidence that they may have fungicidal properties.

Signals and regulators

Plants produce hormones and other growth regulators which act to signal a physiological response in their tissues. They also produce compounds such as phytochrome
Phytochrome
Phytochrome is a photoreceptor, a pigment that plants use to detect light. It is sensitive to light in the red and far-red region of the visible spectrum. Many flowering plants use it to regulate the time of flowering based on the length of day and night and to set circadian rhythms...

 that are sensitive to light and which serve to trigger growth or development in response to environmental signals.

Plant hormones

Plant hormone
Plant hormone
Plant hormones are chemicals that regulate plant growth, which, in the UK, are termed 'plant growth substances'. Plant hormones are signal molecules produced within the plant, and occur in extremely low concentrations. Hormones regulate cellular processes in targeted cells locally and, when moved...

s, known as plant growth regulators (PGRs) or phytohormones, are chemicals that regulate a plant's growth. According to a standard animal definition, hormone
Hormone
A hormone is a chemical released by a cell or a gland in one part of the body that sends out messages that affect cells in other parts of the organism. Only a small amount of hormone is required to alter cell metabolism. In essence, it is a chemical messenger that transports a signal from one...

s are signal molecules produced at specific locations, that occur in very low concentrations, and cause altered processes in target cells at other locations. Unlike animals, plants lack specific hormone-producing tissues or organs. Plant hormones are often not transported to other parts of the plant and production is not limited to specific locations.

Plant hormones are chemicals that in small amounts promote and influence the growth
Cell growth
The term cell growth is used in the contexts of cell development and cell division . When used in the context of cell division, it refers to growth of cell populations, where one cell grows and divides to produce two "daughter cells"...

, development
Morphogenesis
Morphogenesis , is the biological process that causes an organism to develop its shape...

 and differentiation
Cellular differentiation
In developmental biology, cellular differentiation is the process by which a less specialized cell becomes a more specialized cell type. Differentiation occurs numerous times during the development of a multicellular organism as the organism changes from a simple zygote to a complex system of...

 of cells and tissues. Hormones are vital to plant growth; affecting processes in plants from flowering to seed
Seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...

 development, dormancy
Dormancy
Dormancy is a period in an organism's life cycle when growth, development, and physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be closely associated with environmental conditions...

, and germination
Germination
Germination is the process in which a plant or fungus emerges from a seed or spore, respectively, and begins growth. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However the growth of a sporeling from a spore, for example the...

. They regulate which tissues grow upwards and which grow downwards, leaf formation and stem growth, fruit development and ripening, as well as leaf abscission
Abscission
Abscission is a term used in several areas of biology. In plant sciences it most commonly refers to the process by which a plant drops one or more of its parts, such as a leaf, fruit, flower or seed...

 and even plant death.

The most important plant hormones are abscissic acid (ABA), auxin
Auxin
Auxins are a class of plant hormones with some morphogen-like characteristics. Auxins have a cardinal role in coordination of many growth and behavioral processes in the plant's life cycle and are essential for plant body development. Auxins and their role in plant growth were first described by...

s, ethylene
Ethylene
Ethylene is a gaseous organic compound with the formula . It is the simplest alkene . Because it contains a carbon-carbon double bond, ethylene is classified as an unsaturated hydrocarbon. Ethylene is widely used in industry and is also a plant hormone...

, gibberellin
Gibberellin
Gibberellins are plant hormones that regulate growth and influence various developmental processes, including stem elongation, germination, dormancy, flowering, sex expression, enzyme induction, and leaf and fruit senescence....

s, and cytokinin
Cytokinin
Cytokinins are a class of plant growth substances that promote cell division, or cytokinesis, in plant roots and shoots. They are involved primarily in cell growth and differentiation, but also affect apical dominance, axillary bud growth, and leaf senescence...

s, though there are many other substances that serve to regulate plant physiology.

Photomorphogenesis

While most people know that light
Light
Light or visible light is electromagnetic radiation that is visible to the human eye, and is responsible for the sense of sight. Visible light has wavelength in a range from about 380 nanometres to about 740 nm, with a frequency range of about 405 THz to 790 THz...

 is important for photosynthesis in plants, few realize that plant sensitivity to light plays a role in the control of plant structural development (morphogenesis
Morphogenesis
Morphogenesis , is the biological process that causes an organism to develop its shape...

). The use of light to control structural development is called photomorphogenesis
Photomorphogenesis
In developmental biology, photomorphogenesis is light-mediated development. The photomorphogenesis of plants is often studied by using tightly-frequency-controlled light sources to grow the plants.-Germination:...

, and is dependent upon the presence of specialized photoreceptors, which are chemical pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...

s capable of absorbing specific wavelength
Wavelength
In physics, the wavelength of a sinusoidal wave is the spatial period of the wave—the distance over which the wave's shape repeats.It is usually determined by considering the distance between consecutive corresponding points of the same phase, such as crests, troughs, or zero crossings, and is a...

s of light.

Plants use four kinds of photoreceptors: phytochrome
Phytochrome
Phytochrome is a photoreceptor, a pigment that plants use to detect light. It is sensitive to light in the red and far-red region of the visible spectrum. Many flowering plants use it to regulate the time of flowering based on the length of day and night and to set circadian rhythms...

, cryptochrome
Cryptochrome
Cryptochromes are a class of blue light-sensitive flavoproteins found in plants and animals. Cryptochromes are involved in the circadian rhythms of plants and animals, and in the sensing of magnetic fields in a number of species...

, a UV-B photoreceptor, and protochlorophyllide a. The first two of these, phytochrome and cryptochrome, are photoreceptor protein
Photoreceptor protein
Photoreceptors are light-sensitive proteins involved in the sensing and response to light in a variety of organisms. Some examples are rhodopsin in the photoreceptor cells of the vertebrate retina, phytochrome in plants, and bacteriorhodopsin and bacteriophytochromes in some bacteria...

s, complex molecular structures formed by joining a protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

 with a light-sensitive pigment. Cryptochrome is also known as the UV-A photoreceptor, because it absorbs ultraviolet
Ultraviolet
Ultraviolet light is electromagnetic radiation with a wavelength shorter than that of visible light, but longer than X-rays, in the range 10 nm to 400 nm, and energies from 3 eV to 124 eV...

 light in the long wave "A" region. The UV-B receptor is one or more compounds not yet identified with certainty, though some evidence suggests carotene
Carotene
The term carotene is used for several related unsaturated hydrocarbon substances having the formula C40Hx, which are synthesized by plants but cannot be made by animals. Carotene is an orange photosynthetic pigment important for photosynthesis. Carotenes are all coloured to the human eye...

 or riboflavin
Riboflavin
Riboflavin, also known as vitamin B2 or additive E101, is an easily absorbed micronutrient with a key role in maintaining health in humans and animals. It is the central component of the cofactors FAD and FMN, and is therefore required by all flavoproteins. As such, vitamin B2 is required for a...

 as candidates. Protochlorophyllide a, as its name suggests, is a chemical precursor of chlorophyll
Chlorophyll
Chlorophyll is a green pigment found in almost all plants, algae, and cyanobacteria. Its name is derived from the Greek words χλωρος, chloros and φύλλον, phyllon . Chlorophyll is an extremely important biomolecule, critical in photosynthesis, which allows plants to obtain energy from light...

.

The most studied of the photoreceptors in plants is phytochrome
Phytochrome
Phytochrome is a photoreceptor, a pigment that plants use to detect light. It is sensitive to light in the red and far-red region of the visible spectrum. Many flowering plants use it to regulate the time of flowering based on the length of day and night and to set circadian rhythms...

. It is sensitive to light in the red
Red
Red is any of a number of similar colors evoked by light consisting predominantly of the longest wavelengths of light discernible by the human eye, in the wavelength range of roughly 630–740 nm. Longer wavelengths than this are called infrared , and cannot be seen by the naked eye...

 and far-red
Far-red
Far-red light is light at the extreme red end of the visible spectrum, between red and infra-red light. Usually regarded as the region between 700 and 800 nm wavelength, it is dimly visible to some eyes. It is reflected or transmitted by plants because of the absorbance spectrum of...

 region of the visible spectrum
Visible spectrum
The visible spectrum is the portion of the electromagnetic spectrum that is visible to the human eye. Electromagnetic radiation in this range of wavelengths is called visible light or simply light. A typical human eye will respond to wavelengths from about 390 to 750 nm. In terms of...

. Many flowering plants use it to regulate the time of flower
Flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...

ing based on the length of day and night (photoperiodism
Photoperiodism
Photoperiodism is the physiological reaction of organisms to the length of day or night. It occurs in plants and animals.Photoperiodism can also be defined as the developmental responses of plants to the relative lengths of the light and dark periods...

) and to set circadian rhythms. It also regulates other responses including the germination of seeds, elongation of seedlings, the size, shape and number of leaves, the synthesis of chlorophyll, and the straightening of the epicotyl
Epicotyl
In plant physiology, the epicotyl is the embryonic shoot above the cotyledons. In most plants the epicotyl will eventually develop into the leaves of the plant. In dicots, the hypocotyl is what appears to be the base stem under the spent withered cotyledons, and the shoot just above that is the...

 or hypocotyl
Hypocotyl
The hypocotyl is the stem of a germinating seedling, found below the cotyledons and above the radicle .-Dicots:...

 hook of dicot seedlings.

Photoperiodism

Many flowering plant
Flowering plant
The flowering plants , also known as Angiospermae or Magnoliophyta, are the most diverse group of land plants. Angiosperms are seed-producing plants like the gymnosperms and can be distinguished from the gymnosperms by a series of synapomorphies...

s use the pigment phytochrome to sense seasonal changes in day
Day
A day is a unit of time, commonly defined as an interval equal to 24 hours. It also can mean that portion of the full day during which a location is illuminated by the light of the sun...

 length, which they take as signals to flower. This sensitivity to day length is termed photoperiodism
Photoperiodism
Photoperiodism is the physiological reaction of organisms to the length of day or night. It occurs in plants and animals.Photoperiodism can also be defined as the developmental responses of plants to the relative lengths of the light and dark periods...

. Broadly speaking, flowering plants can be classified as long day plants, short day plants, or day neutral plants, depending on their particular response to changes in day length. Long day plants require a certain minimum length of daylight to initiate flowering, so these plants flower in the spring or summer. Conversely, short day plants flower when the length of daylight falls below a certain critical level. Day neutral plants do not initiate flowering based on photoperiodism, though some may use temperature sensitivity (vernalization
Vernalization
Vernalization is the acquisition of a plant's ability to flower or germinate in the spring by exposure to the prolonged cold of winter...

) instead.

Although a short day plant cannot flower during the long days of summer, it is not actually the period of light exposure that limits flowering. Rather, a short day plant requires a minimal length of uninterrupted darkness in each 24 hour period (a short daylength) before floral development can begin. It has been determined experimentally that a short day plant (long night) does not flower if a flash of phytochrome activating light is used on the plant during the night.

Plants make use of the phytochrome system to sense day length or photoperiod. This fact is utilized by florists and greenhouse
Greenhouse
A greenhouse is a building in which plants are grown. These structures range in size from small sheds to very large buildings...

 gardeners to control and even induce flowering out of season, such as the Poinsettia
Poinsettia
Euphorbia pulcherrima, commonly known as Zack Wood or noche buena, is a species of flowering plant indigenous to Mexico and Central America. The name "poinsettia" is after Joel Roberts Poinsett, the first United States Minister to Mexico, who introduced the plant into the US in 1825...

.

Environmental physiology

Paradoxically, the subdiscipline of environmental physiology is on the one hand a recent field of study in plant ecology and on the other hand one of the oldest. Environmental physiology is the preferred name of the subdiscipline among plant physiologists, but it goes by a number of other names in the applied sciences. It is roughly synonymous with ecophysiology
Ecophysiology
Ecophysiology or environmental physiology is a biological discipline which studies the adaptation of organism's physiology to environmental conditions...

, crop ecology, horticulture
Horticulture
Horticulture is the industry and science of plant cultivation including the process of preparing soil for the planting of seeds, tubers, or cuttings. Horticulturists work and conduct research in the disciplines of plant propagation and cultivation, crop production, plant breeding and genetic...

 and agronomy
Agronomy
Agronomy is the science and technology of producing and using plants for food, fuel, feed, fiber, and reclamation. Agronomy encompasses work in the areas of plant genetics, plant physiology, meteorology, and soil science. Agronomy is the application of a combination of sciences like biology,...

. The particular name applied to the subdiscipline is specific to the viewpoint and goals of research. Whatever name is applied, it deals with the ways in which plants respond to their environment and so overlaps with the field of ecology
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...

.

Environmental physiologists examine plant response to physical factors such as radiation
Radiation
In physics, radiation is a process in which energetic particles or energetic waves travel through a medium or space. There are two distinct types of radiation; ionizing and non-ionizing...

 (including light
Light
Light or visible light is electromagnetic radiation that is visible to the human eye, and is responsible for the sense of sight. Visible light has wavelength in a range from about 380 nanometres to about 740 nm, with a frequency range of about 405 THz to 790 THz...

 and ultraviolet
Ultraviolet
Ultraviolet light is electromagnetic radiation with a wavelength shorter than that of visible light, but longer than X-rays, in the range 10 nm to 400 nm, and energies from 3 eV to 124 eV...

 radiation), temperature
Temperature
Temperature is a physical property of matter that quantitatively expresses the common notions of hot and cold. Objects of low temperature are cold, while various degrees of higher temperatures are referred to as warm or hot...

, fire
Fire
Fire is the rapid oxidation of a material in the chemical process of combustion, releasing heat, light, and various reaction products. Slower oxidative processes like rusting or digestion are not included by this definition....

, and wind
Wind
Wind is the flow of gases on a large scale. On Earth, wind consists of the bulk movement of air. In outer space, solar wind is the movement of gases or charged particles from the sun through space, while planetary wind is the outgassing of light chemical elements from a planet's atmosphere into space...

. Of particular importance are water
Water
Water is a chemical substance with the chemical formula H2O. A water molecule contains one oxygen and two hydrogen atoms connected by covalent bonds. Water is a liquid at ambient conditions, but it often co-exists on Earth with its solid state, ice, and gaseous state . Water also exists in a...

 relations (which can be measured with the Pressure bomb
Pressure bomb
A pressure bomb or pressure chamber or Scholander bomb is an instrument with which it is possible to measure the approximate water potential of plant tissues. A leaf attached to a stem is placed inside a sealed chamber and pressurised gas is added to the chamber slowly...

) and the stress of drought
Drought
A drought is an extended period of months or years when a region notes a deficiency in its water supply. Generally, this occurs when a region receives consistently below average precipitation. It can have a substantial impact on the ecosystem and agriculture of the affected region...

 or inundation
Flood
A flood is an overflow of an expanse of water that submerges land. The EU Floods directive defines a flood as a temporary covering by water of land not normally covered by water...

, exchange of gases with the atmosphere
Atmosphere
An atmosphere is a layer of gases that may surround a material body of sufficient mass, and that is held in place by the gravity of the body. An atmosphere may be retained for a longer duration, if the gravity is high and the atmosphere's temperature is low...

, as well as the cycling of nutrients such as nitrogen
Nitrogen
Nitrogen is a chemical element that has the symbol N, atomic number of 7 and atomic mass 14.00674 u. Elemental nitrogen is a colorless, odorless, tasteless, and mostly inert diatomic gas at standard conditions, constituting 78.08% by volume of Earth's atmosphere...

 and carbon
Carbon
Carbon is the chemical element with symbol C and atomic number 6. As a member of group 14 on the periodic table, it is nonmetallic and tetravalent—making four electrons available to form covalent chemical bonds...

.

Environmental physiologists also examine plant response to biological factors. This includes not only negative interactions, such as competition
Competition (biology)
Competition is an interaction between organisms or species, in which the fitness of one is lowered by the presence of another. Limited supply of at least one resource used by both is required. Competition both within and between species is an important topic in ecology, especially community ecology...

, herbivory, disease
Disease
A disease is an abnormal condition affecting the body of an organism. It is often construed to be a medical condition associated with specific symptoms and signs. It may be caused by external factors, such as infectious disease, or it may be caused by internal dysfunctions, such as autoimmune...

 and parasitism
Parasitism
Parasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host . These are now called macroparasites...

, but also positive interactions, such as mutualism and pollination
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...

.

Tropisms and nastic movements

Plants may respond both to directional and nondirectional stimuli
Stimulus (physiology)
In physiology, a stimulus is a detectable change in the internal or external environment. The ability of an organism or organ to respond to external stimuli is called sensitivity....

. A response to a directional stimulus, such as gravity or sunlight
Light
Light or visible light is electromagnetic radiation that is visible to the human eye, and is responsible for the sense of sight. Visible light has wavelength in a range from about 380 nanometres to about 740 nm, with a frequency range of about 405 THz to 790 THz...

, is called a tropism. A response to a nondirectional stimulus, such as temperature
Temperature
Temperature is a physical property of matter that quantitatively expresses the common notions of hot and cold. Objects of low temperature are cold, while various degrees of higher temperatures are referred to as warm or hot...

 or humidity
Humidity
Humidity is a term for the amount of water vapor in the air, and can refer to any one of several measurements of humidity. Formally, humid air is not "moist air" but a mixture of water vapor and other constituents of air, and humidity is defined in terms of the water content of this mixture,...

, is a nastic movement.

Tropism
Tropism
A tropism is a biological phenomenon, indicating growth or turning movement of a biological organism, usually a plant, in response to an environmental stimulus. In tropisms, this response is dependent on the direction of the stimulus...

s in plants are the result of differential cell
Cell (biology)
The cell is the basic structural and functional unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often called the building block of life. The Alberts text discusses how the "cellular building blocks" move to shape developing embryos....

 growth, in which the cells on one side of the plant elongate more than those on the other side, causing the part to bend toward the side with less growth. Among the common tropisms seen in plants is phototropism
Phototropism
Phototropism is directional growth in which the direction of growth is determined by the direction of the light source. In other words, it is the growth and response to a light stimulus. Phototropism is most often observed in plants, but can also occur in other organisms such as fungi...

, the bending of the plant toward a source of light. Phototropism allows the plant to maximize light exposure in plants which require additional light for photosynthesis, or to minimize it in plants subjected to intense light and heat. Geotropism allows the roots of a plant to determine the direction of gravity and grow downwards. Tropisms generally result from an interaction between the environment and production of one or more plant hormones.

In contrast to tropisms, nastic movements result from changes in turgor pressure
Turgor pressure
Turgor Pressure or turgidity is the main pressure of the cell contents against the cell wall in plant cells and bacteria cells, determined by the water content of the vacuole, resulting from osmotic pressure, i.e...

 within plant tissues, and may occur rapidly. A familiar example is thigmonasty
Thigmonasty
Thigmonasty or seismonasty is the nastic response of a plant or fungus to touch or vibration. It differs from thigmotropism in that it is independent of the direction of the stimulus. For example, tendrils from a climbing plant are thigmotropic because they twine around any support they touch. ...

 (response to touch) in the Venus fly trap, a carnivorous plant
Carnivorous plant
Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans, typically insects and other arthropods. Carnivorous plants appear adapted to grow in places where the soil is thin or poor in nutrients, especially nitrogen, such as acidic...

. The traps consist of modified leaf blades which bear sensitive trigger hairs. When the hairs are touched by an insect or other animal, the leaf folds shut. This mechanism allows the plant to trap and digest small insects for additional nutrients. Although the trap is rapidly shut by changes in internal cell pressures, the leaf must grow slowly to reset for a second opportunity to trap insects.

Plant disease

Economically, one of the most important areas of research in environmental physiology is that of phytopathology
Phytopathology
Plant pathology is the scientific study of plant diseases caused by pathogens and environmental conditions . Organisms that cause infectious disease include fungi, oomycetes, bacteria, viruses, viroids, virus-like organisms, phytoplasmas, protozoa, nematodes and parasitic plants...

, the study of disease
Disease
A disease is an abnormal condition affecting the body of an organism. It is often construed to be a medical condition associated with specific symptoms and signs. It may be caused by external factors, such as infectious disease, or it may be caused by internal dysfunctions, such as autoimmune...

s in plants and the manner in which plants resist or cope with infection. Plant are susceptible to the same kinds of disease organisms as animals, including virus
Virus
A virus is a small infectious agent that can replicate only inside the living cells of organisms. Viruses infect all types of organisms, from animals and plants to bacteria and archaea...

es, bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...

, and fungi, as well as physical invasion by insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s and roundworms.

Because the biology of plants differs from animals, their symptoms and responses are quite different. In some cases, a plant can simply shed infected leaves or flowers to prevent to spread of disease, in a process called abscission. Most animals do not have this option as a means of controlling disease. Plant diseases organisms themselves also differ from those causing disease in animals because plants cannot usually spread infection through casual physical contact. Plant pathogen
Pathogen
A pathogen gignomai "I give birth to") or infectious agent — colloquially, a germ — is a microbe or microorganism such as a virus, bacterium, prion, or fungus that causes disease in its animal or plant host...

s tend to spread via spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

s or are carried by animal vectors.

One of the most important advances in the control of plant disease was the discovery of Bordeaux mixture
Bordeaux mixture
Bordeaux mixture is a mixture of copper sulfate and slaked lime used as a fungicide in vineyards. It is used mainly to control garden, vineyard, nursery and farm infestations of fungi, primarily downy mildew which can result from infections of Plasmopara viticola. It was invented in the Bordeaux...

 in the nineteenth century. The mixture is the first known fungicide
Fungicide
Fungicides are chemical compounds or biological organisms used to kill or inhibit fungi or fungal spores. Fungi can cause serious damage in agriculture, resulting in critical losses of yield, quality and profit. Fungicides are used both in agriculture and to fight fungal infections in animals...

 and is a combination of copper sulfate and lime
Lime (mineral)
Lime is a general term for calcium-containing inorganic materials, in which carbonates, oxides and hydroxides predominate. Strictly speaking, lime is calcium oxide or calcium hydroxide. It is also the name for a single mineral of the CaO composition, occurring very rarely...

. Application of the mixture served to inhibit the growth of downy mildew
Downy mildew
Downy mildew refers to any of several types of oomycete microbes that are obligate parasites of plants. Downy mildews exclusively belong to Peronosporaceae. In commercial agriculture, they are a particular problem for growers of crucifers, grapes and vegetables that grow on vines...

 that threatened to seriously damage the French
France
The French Republic , The French Republic , The French Republic , (commonly known as France , is a unitary semi-presidential republic in Western Europe with several overseas territories and islands located on other continents and in the Indian, Pacific, and Atlantic oceans. Metropolitan France...

 wine
Wine
Wine is an alcoholic beverage, made of fermented fruit juice, usually from grapes. The natural chemical balance of grapes lets them ferment without the addition of sugars, acids, enzymes, or other nutrients. Grape wine is produced by fermenting crushed grapes using various types of yeast. Yeast...

 industry.

Early history

Sir Francis Bacon
Francis Bacon
Francis Bacon, 1st Viscount St Albans, KC was an English philosopher, statesman, scientist, lawyer, jurist, author and pioneer of the scientific method. He served both as Attorney General and Lord Chancellor of England...

 published one of the first plant physiology experiments in 1627 in the book, Sylva Sylvarum. Bacon grew several terrestrial plants, including a rose, in water and concluded that soil was only needed to keep the plant upright. Jan Baptist van Helmont
Jan Baptist van Helmont
Jan Baptist van Helmont was an early modern period Flemish chemist, physiologist, and physician. He worked during the years just after Paracelsus and iatrochemistry, and is sometimes considered to be "the founder of pneumatic chemistry"...

 published what is considered the first quantitative experiment in plant physiology in 1648. He grew a willow tree for five years in a pot containing 200 pounds of oven-dry soil. The soil lost just two ounces of dry weight and van Helmont concluded that plants get all their weight from water, not soil. In 1699, John Woodward
John Woodward (naturalist)
John Woodward was an English naturalist, antiquarian and geologist, and founder by bequest of the Woodwardian Professorship of Geology at Cambridge University...

 published experiments on growth of spearmint
Spearmint
Mentha spicata syn. M. cordifolia is a species of mint native to much of Europe and southwest Asia, though its exact natural range is uncertain due to extensive early cultivation. It grows in wet soils...

 in different sources of water. He found that plants grew much better in water with soil added than in distilled water.

Stephen Hales
Stephen Hales
Stephen Hales, FRS was an English physiologist, chemist and inventor.Hales studied the role of air and water in the maintenance of both plant and animal life. He gave accurate accounts of the movements of water in plants, and demonstrated that plants absorb air...

 is considered the Father of Plant Physiology for the many experiments in the 1727 book; though
Julius von Sachs
Julius von Sachs
Julius von Sachs was a German botanist from Breslau, Prussian Silesia.At an early age he showed a taste for natural history, becoming acquainted with the Breslau physiologist Jan Evangelista Purkyně. In 1851 he began studying at Charles University in Prague...

 unified the pieces of plant physiology and put them together as a discipline. His Lehrbuch der Botanik was the plant physiology bible of its time.

Researchers discovered in the 1800s that plants absorb essential mineral nutrients as inorganic ions in water. In natural conditions, soil acts as a mineral nutrient reservoir but the soil itself is not essential to plant growth. When the mineral nutrients in the soil are dissolved in water, plant roots absorb nutrients readily, soil is no longer required for the plant to thrive. This observation is the basis for hydroponics
Hydroponics
Hydroponics is a method of growing plants using mineral nutrient solutions, in water, without soil. Terrestrial plants may be grown with their roots in the mineral nutrient solution only or in an inert medium, such as perlite, gravel, mineral wool, or coconut husk.Researchers discovered in the 18th...

, the growing of plants in a water solution rather than soil, which has become a standard technique in biological research, teaching lab exercises, crop production and as a hobby.

Current research

One of the leading journals in the field is Plant Physiology
Plant Physiology (journal)
Plant Physiology is a monthly peer-reviewed scientific journal that covers research on physiology, biochemistry, cellular and molecular biology, genetics, biophysics, and environmental biology of plants. The journal has been published since 1926 by the American Society of Plant Biologists. It is a...

, started in 1926. All its back issues are available online for free.http://www.plantphysiol.org/contents-by-date.0.shtml Many other journals often carry plant physiology articles, including Physiologia Plantarum, Journal of Experimental Botany, American Journal of Botany
American Journal of Botany
The American Journal of Botany is a peer-reviewed scientific journal which includes research papers on all aspects of plant biology. It is published by the Botanical Society of America on a monthly basis since 1914....

, Annals of Botany, Journal of Plant Nutrition and Proceedings of the National Academy of Sciences
Proceedings of the National Academy of Sciences
The Proceedings of the National Academy of Sciences of the United States of America, usually referred to as PNAS, is the official journal of the United States National Academy of Sciences...

.

Food production

In horticulture
Horticulture
Horticulture is the industry and science of plant cultivation including the process of preparing soil for the planting of seeds, tubers, or cuttings. Horticulturists work and conduct research in the disciplines of plant propagation and cultivation, crop production, plant breeding and genetic...

 and agriculture
Agriculture
Agriculture is the cultivation of animals, plants, fungi and other life forms for food, fiber, and other products used to sustain life. Agriculture was the key implement in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that nurtured the...

 along with food science
Food science
Food science is a study concerned with all technical aspects of foods, beginning with harvesting or slaughtering, and ending with its cooking and consumption, an ideology commonly referred to as "from field to fork"...

, plant physiology is an important topic relating to fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...

s, vegetable
Vegetable
The noun vegetable usually means an edible plant or part of a plant other than a sweet fruit or seed. This typically means the leaf, stem, or root of a plant....

s, and other consumable parts of plants. Topics studied include: climatic requirements, fruit drop, nutrition, ripening
Ripening
Ripening is a process in fruits that causes them to become more palatable. In general, a fruit becomes sweeter, less green, and softer as it ripens. Even though the acidity of fruit increases as it ripens, the higher acidity level does not make the fruit seem tarter, which can lead to the...

, fruit set. The production of food crops also hinges on the study of plant physiology covering such topics as Optimal planting and harvesting times and post harvest storage of plant products for human consumption and the production of secondary products like drugs and cosmetics.

See also

  • Biomechanics
    Biomechanics
    Biomechanics is the application of mechanical principles to biological systems, such as humans, animals, plants, organs, and cells. Perhaps one of the best definitions was provided by Herbert Hatze in 1974: "Biomechanics is the study of the structure and function of biological systems by means of...

  • Phytochemistry
    Phytochemistry
    Phytochemistry is in the strict sense of the word the study of phytochemicals. These are chemicals derived from plants. In a narrower sense the terms are often used to describe the large number of secondary metabolic compounds found in plants. Many of these are known to provide protection against...

  • Plant anatomy
    Plant anatomy
    Plant anatomy or phytotomy is the general term for the study of the internal structure of plants. While originally it included plant morphology, which is the description of the physical form and external structure of plants, since the mid-20th century the investigations of plant anatomy are...

  • Plant morphology
    Plant morphology
    Plant morphology or phytomorphology is the study of the physical form and external structure of plants. This is usually considered distinct from plant anatomy, which is the study of the internal structure of plants, especially at the microscopic level...




Further reading

  • Salisbury, Frank B. & Ross, Cleon W. (1992). Plant physiology, 4th, Belmont, California: Wadsworth Publishing. ISBN 0-534-15162-0.

External links


Scientific journals

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