Plant physiology

Plant physiology is a subdiscipline of botany concerned with the functioning, or physiology, of plants.[1] Closely related fields include plant morphology (structure of plants), plant ecology (interactions with the environment), phytochemistry (biochemistry of plants), cell biology, genetics, biophysics and molecular biology.

Fundamental processes such as photosynthesis, respiration, plant nutrition, plant hormone functions, tropisms, nastic movements, photoperiodism, photomorphogenesis, circadian rhythms, environmental stress physiology, seed germination, dormancy and stomata function and transpiration, both parts of plant water relations, are studied by plant physiologists.

Kiemtafel (germination table)
A germination rate experiment


The field of plant physiology includes the study of all the internal activities of plants—those chemical and physical processes associated with life as they occur in plants. This includes study at many levels of scale of size and time. At the smallest scale are molecular interactions of photosynthesis and internal diffusion of water, minerals, and nutrients. At the largest scale are the processes of plant development, seasonality, dormancy, and reproductive control. Major subdisciplines of plant physiology include phytochemistry (the study of the biochemistry of plants) and phytopathology (the study of disease in plants). The scope of plant physiology as a discipline may be divided into several major areas of research.

Plant physiology
Five key areas of study within plant physiology.

First, the study of phytochemistry (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 requires a large array of pigments, enzymes, and other compounds to function. Because they cannot move, plants must also defend themselves chemically from herbivores, pathogens and competition from other plants. They do this by producing toxins 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 pollinators or herbivores to spread ripe seeds.

Secondly, plant physiology includes the study of biological and chemical processes of individual plant cells. Plant cells have a number of features that distinguish them from cells of animals, 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 which restricts the shape of plant cells and thereby limits the flexibility and mobility of plants. Plant cells also contain chlorophyll, a chemical compound that interacts with light 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, and organs within a plant. Different cells and tissues are physically and chemically specialized to perform different functions. Roots and rhizoids function to anchor the plant and acquire minerals in the soil. Leaves 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, 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 which are produced in one part of the plant to signal cells in another part of the plant to respond. Many flowering plants bloom at the appropriate time because of light-sensitive compounds that respond to the length of the night, a phenomenon known as photoperiodism. The ripening of fruit and loss of leaves in the winter are controlled in part by the production of the gas ethylene by the plant.

Finally, plant physiology includes the study of plant response to environmental conditions and their variation, 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

Latex dripping
Latex being collected from a tapped rubber tree.

The chemical elements of which plants are constructed—principally carbon, oxygen, hydrogen, nitrogen, phosphorus, sulfur, etc.—are the same as for all other life forms animals, fungi, bacteria and even viruses. Only the details of the molecules into which they are assembled differs.

Despite this underlying similarity, plants produce a vast array of chemical compounds with unique properties which they use to cope with their environment. Pigments are used by plants to absorb or detect light, and are extracted by humans for use in dyes. Other plant products may be used for the manufacture of commercially important rubber or biofuel. Perhaps the most celebrated compounds from plants are those with pharmacological activity, such as salicylic acid from which aspirin is made, morphine, and digoxin. Drug companies spend billions of dollars each year researching plant compounds for potential medicinal benefits.

Constituent elements

Plants require some nutrients, such as carbon and nitrogen, in large quantities to survive. Some 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 micronutrients, are required only in trace amounts for plants to remain healthy. Such micronutrients are usually absorbed as ions dissolved in water taken from the soil, though carnivorous plants acquire some of their micronutrients from captured prey.

The following tables list element nutrients essential to plants. Uses within plants are generalized.

Macronutrients – necessary in large quantities
Element Form of uptake Notes
Nitrogen NO3, NH4+ Nucleic acids, proteins, hormones, etc.
Oxygen O2 H2O Cellulose, starch, other organic compounds
Carbon CO2 Cellulose, starch, other organic compounds
Hydrogen H2O Cellulose, starch, other organic compounds
Potassium K+ Cofactor in protein synthesis, water balance, etc.
Calcium Ca2+ Membrane synthesis and stabilization
Magnesium Mg2+ Element essential for chlorophyll
Phosphorus H2PO4 Nucleic acids, phospholipids, ATP
Sulfur SO42− Constituent of proteins
Micronutrients – necessary in small quantities
Element Form of uptake Notes
Chlorine Cl Photosystem II and stomata function
Iron Fe2+, Fe3+ Chlorophyll formation and nitrogen fixation
Boron HBO3 Crosslinking pectin
Manganese Mn2+ Activity of some enzymes and photosystem II
Zinc Zn2+ Involved in the synthesis of enzymes and chlorophyll
Copper Cu+ Enzymes for lignin synthesis
Molybdenum MoO42− Nitrogen fixation, reduction of nitrates
Nickel Ni2+ Enzymatic cofactor in the metabolism of nitrogen compounds


Space-filling model of the chlorophyll molecule.
Orange violet pansies
Anthocyanin gives these pansies their dark purple pigmentation.

Among the most important molecules for plant function are the pigments. Plant pigments include a variety of different kinds of molecules, including porphyrins, carotenoids, and anthocyanins. All biological pigments selectively absorb certain wavelengths of light while reflecting others. The light that is absorbed may be used by the plant to power chemical reactions, while the reflected wavelengths of light determine the color the pigment appears to the eye.

Chlorophyll is the primary pigment in plants; it is a porphyrin that absorbs red and blue wavelengths of light while reflecting green. 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. Kelps, diatoms, and other photosynthetic heterokonts contain chlorophyll c instead of b, red algae possess chlorophyll a. All chlorophylls serve as the primary means plants use to intercept light to fuel photosynthesis.

Carotenoids are red, orange, or yellow tetraterpenoids. They function as accessory pigments in plants, helping to fuel photosynthesis by gathering wavelengths of light not readily absorbed by chlorophyll. The most familiar carotenoids are carotene (an orange pigment found in carrots), lutein (a yellow pigment found in fruits and vegetables), and lycopene (the red pigment responsible for the color of tomatoes). Carotenoids have been shown to act as antioxidants and to promote healthy eyesight in humans.

Anthocyanins (literally "flower blue") are water-soluble flavonoid pigments that appear red to blue, according to pH. They occur in all tissues of higher plants, providing color in leaves, stems, roots, flowers, and fruits, though not always in sufficient quantities to be noticeable. Anthocyanins are most visible in the petals of flowers, where they may make up as much as 30% of the dry weight of the tissue.[2] They are also responsible for the purple color seen on the underside of tropical shade plants such as Tradescantia zebrina. 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

Betalains are red or yellow pigments. Like anthocyanins they are water-soluble, but unlike anthocyanins they are indole-derived compounds synthesized from tyrosine. This class of pigments is found only in the Caryophyllales (including cactus and amaranth), and never co-occur in plants with anthocyanins. Betalains are responsible for the deep red color of beets, 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.[3]

Signals and regulators

A mutation that stops Arabidopsis thaliana responding to auxin causes abnormal growth (right)

Plants produce hormones and other growth regulators which act to signal a physiological response in their tissues. They also produce compounds such as phytochrome that are sensitive to light and which serve to trigger growth or development in response to environmental signals.

Plant hormones

Plant hormones, known as plant growth regulators (PGRs) or phytohormones, are chemicals that regulate a plant's growth. According to a standard animal definition, hormones 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, development and differentiation of cells and tissues. Hormones are vital to plant growth; affecting processes in plants from flowering to seed development, dormancy, and germination. They regulate which tissues grow upwards and which grow downwards, leaf formation and stem growth, fruit development and ripening, as well as leaf abscission and even plant death.

The most important plant hormones are abscissic acid (ABA), auxins, ethylene, gibberellins, and cytokinins, though there are many other substances that serve to regulate plant physiology.


While most people know that light is important for photosynthesis in plants, few realize that plant sensitivity to light plays a role in the control of plant structural development (morphogenesis). The use of light to control structural development is called photomorphogenesis, and is dependent upon the presence of specialized photoreceptors, which are chemical pigments capable of absorbing specific wavelengths of light.

Plants use four kinds of photoreceptors:[1] phytochrome, cryptochrome, a UV-B photoreceptor, and protochlorophyllide a. The first two of these, phytochrome and cryptochrome, are photoreceptor proteins, complex molecular structures formed by joining a protein with a light-sensitive pigment. Cryptochrome is also known as the UV-A photoreceptor, because it absorbs ultraviolet 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 or riboflavin as candidates.[4] Protochlorophyllide a, as its name suggests, is a chemical precursor of chlorophyll.

The most studied of the photoreceptors in plants is phytochrome. 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 (photoperiodism) 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 or hypocotyl hook of dicot seedlings.


Poinsettia 2
The poinsettia is a short-day plant, requiring two months of long nights prior to blooming.

Many flowering plants use the pigment phytochrome to sense seasonal changes in day length, which they take as signals to flower. This sensitivity to day length is termed photoperiodism. 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 starts 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) 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 gardeners to control and even induce flowering out of season, such as the Poinsettia.

Environmental physiology

Arabidopsis thaliana
Phototropism in Arabidopsis thaliana is regulated by blue to UV light.[5]

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.[1] 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, crop ecology, horticulture and agronomy. 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.

Environmental physiologists examine plant response to physical factors such as radiation (including light and ultraviolet radiation), temperature, fire, and wind. Of particular importance are water relations (which can be measured with the Pressure bomb) and the stress of drought or inundation, exchange of gases with the atmosphere, as well as the cycling of nutrients such as nitrogen and carbon.

Environmental physiologists also examine plant response to biological factors. This includes not only negative interactions, such as competition, herbivory, disease and parasitism, but also positive interactions, such as mutualism and pollination.

Tropisms and nastic movements

Plants may respond both to directional and non-directional stimuli. A response to a directional stimulus, such as gravity or sunlight, is called a tropism. A response to a nondirectional stimulus, such as temperature or humidity, is a nastic movement.

Tropisms in plants are the result of differential cell growth, in which the cells on one side of the plant elongates 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, 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.

Nastic movements results from differential cell growth (e.g. epinasty and hiponasty), or from changes in turgor pressure within plant tissues (e.g., nyctinasty), which may occur rapidly. A familiar example is thigmonasty (response to touch) in the Venus fly trap, a carnivorous plant. 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.[6]

Plant disease

Powdery mildew
Powdery mildew on crop leaves

Economically, one of the most important areas of research in environmental physiology is that of phytopathology, the study of diseases 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 viruses, bacteria, and fungi, as well as physical invasion by insects and roundworms.

Because the biology of plants differs with animals, their symptoms and responses are quite different. In some cases, a plant can simply shed infected leaves or flowers to prevent the 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 pathogens tend to spread via spores or are carried by animal vectors.

One of the most important advances in the control of plant disease was the discovery of Bordeaux mixture in the nineteenth century. The mixture is the first known fungicide and is a combination of copper sulfate and lime. Application of the mixture served to inhibit the growth of downy mildew that threatened to seriously damage the French wine industry.[7]


Early history

Sir Francis Bacon 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 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 published experiments on growth of spearmint in different sources of water. He found that plants grew much better in water with soil added than in distilled water.

Stephen Hales is considered the Father of Plant Physiology for the many experiments in the 1727 book;[8] though Julius von Sachs 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.[9]

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, 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, started in 1926. All its back issues are available online for free.[1] Many other journals often carry plant physiology articles, including Physiologia Plantarum, Journal of Experimental Botany, American Journal of Botany, Annals of Botany, Journal of Plant Nutrition and Proceedings of the National Academy of Sciences.

Economic applications

Food production

In horticulture and agriculture along with food science, plant physiology is an important topic relating to fruits, vegetables, and other consumable parts of plants. Topics studied include: climatic requirements, fruit drop, nutrition, ripening, 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


  1. ^ a b c Frank B. Salisbury; Cleon W. Ross (1992). Plant physiology. Brooks/Cole Pub Co. ISBN 0-534-15162-0.
  2. ^ Trevor Robinson (1963). The organic constituents of higher plants: their chemistry and interrelationships. Cordus Press. p. 183.
  3. ^ Kimler, L. M. (1975). "Betanin, the red beet pigment, as an antifungal agent". Botanical Society of America, Abstracts of papers. 36.
  4. ^ Fosket, Donald E. (1994). Plant Growth and Development: A Molecular Approach. San Diego: Academic Press. pp. 498–509. ISBN 0-12-262430-0.
  5. ^
  6. ^ Adrian Charles Slack; Jane Gate (1980). Carnivorous Plants. Cambridge, Massachusetts : MIT Press. p. 160. ISBN 978-0-262-19186-9.
  7. ^ Kingsley Rowland Stern; Shelley Jansky (1991). Introductory Plant Biology. WCB/McGraw-Hill. p. 309. ISBN 978-0-697-09948-8.
  8. ^ Hales, Stephen. 1727. Vegetable Staticks
  9. ^ Duane Isely (1994). 101 Botanists. Iowa State Press. pp. 216–219. ISBN 978-0-8138-2498-7.

Further reading

  • Lambers, H. (1998). Plant physiological ecology. New York: Springer-Verlag. ISBN 0-387-98326-0.
  • Larcher, W. (2001). Physiological plant ecology (4th ed.). Springer. ISBN 3-540-43516-6.
  • Frank B. Salisbury; Cleon W. Ross (1992). Plant physiology. Brooks/Cole Pub Co. ISBN 0-534-15162-0.
  • Lincoln Taiz, Eduardo Zeiger, Ian Max Møller, Angus Murphy: Fundamentals of Plant Physiology. Sinauer, 2018.

External links

Scientific journals


Agronomy (from Ancient Greek ἀγρός agrós "field" and νόμος nómos "law") is the science and technology of producing and using plants for food, fuel, fiber, and land reclamation. Agronomy has come to encompass work in the areas of plant genetics, plant physiology, meteorology, and soil science. It is the application of a combination of sciences like biology, chemistry, economics, ecology, earth science, and genetics. Agronomists of today are involved with many issues, including producing food, creating healthier food, managing the environmental impact of agriculture, and extracting energy from plants. Agronomists often specialise in areas such as crop rotation, irrigation and drainage, plant breeding, plant physiology, soil classification, soil fertility, weed control, and insect and pest control.


Amyloplasts are a type of plastid, double-enveloped organelles in plant cells that are involved in various biological pathways. Amyloplasts are specifically a type of leucoplast, a subcategory for colorless, non-pigment-containing plastids. Amyloplasts are found in roots and storage tissues and store and synthesize starch for the plant through the polymerization of glucose. Starch synthesis relies on the transportation of carbon from the cytosol, the mechanism by which is currently under debate.Starch synthesis and storage also takes place in chloroplasts, a type of pigmented plastid involved in photosynthesis. Amyloplasts and chloroplasts are closely related, and amyloplasts can turn into chloroplasts; this is for instance observed when potato tubers are exposed to light and turn green.


In botany, a bud is an undeveloped or embryonic shoot and normally occurs in the axil of a leaf or at the tip of a stem. Once formed, a bud may remain for some time in a dormant condition, or it may form a shoot immediately. Buds may be specialized to develop flowers or short shoots, or may have the potential for general shoot development. The term bud is also used in zoology, where it refers to an outgrowth from the body which can develop into a new individual.

Cortex (botany)

A cortex is an outer layer of a stem or root in a plant, lying below the epidermis but outside the vascular bundles.The cortex is composed mostly of large thin-walled parenchyma cells of the ground tissue system and shows little to no structural differentiation. The outer cortical cells often acquire irregularly thickened cell walls, and are called collenchyma cells. Some of the outer cortical cells may contain chloroplasts.Cortex forms layer of cells that constitute cork. It is responsible for the transportation of materials into the central cylinder of the root through diffusion and may also be used for food storage in the form of starch. The innermost layer of the cortex is the endodermis.

Dehiscence (botany)

Dehiscence is the splitting along a built-in line of weakness in a plant structure in order to release its contents, and is common among fruits, anthers and sporangia. Sometimes this involves the complete detachment of a part. Structures that open in this way are said to be dehiscent. Structures that do not open in this way are called indehiscent, and rely on other mechanisms such as decay or predation to release the contents.

A similar process to dehiscence occurs in some flower buds (e.g., Platycodon, Fuchsia), but this is rarely referred to as dehiscence unless circumscissile dehiscence is involved; anthesis is the usual term for the opening of flowers. Dehiscence may or may not involve the loss of a structure through the process of abscission. The lost structures are said to be caducous.


The endosperm is a tissue produced inside the seeds of most of the flowering plants following fertilization. It is triploid in most species. It surrounds the embryo and provides nutrition in the form of starch, though it can also contain oils and protein. This can make endosperm a source of nutrition in animal diet. For example, wheat endosperm is ground into flour for bread (the rest of the grain is included as well in whole wheat flour), while barley endosperm is the main source of sugars for beer production. Other examples of endosperm that forms the bulk of the edible portion are coconut "meat" and coconut "water", and corn. Some plants, such as orchids, lack endosperm in their seeds.


Germination is the process by which an organism grows from a seed or similar structure. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. In addition, the growth of a sporeling from a spore, such as the spores of hyphae from fungal spores, is also germination. Thus, in a general sense, germination can be thought of as anything expanding into greater being from a small existence or germ.

Max Planck Institute of Molecular Plant Physiology

The Max Planck Institute of Molecular Plant Physiology (German: Max-Planck-Institut für molekulare Pflanzenphysiologie) is a German research institute for molecular plant physiology, based in the Golm district of Potsdam, Brandenburg. Founded on January 1, 1994, the MPIMP focuses on the study of the dynamics of plant metabolism and how that relates to the entire plant system. The institution is one of the 80 institutes in the Max Planck Society (Max-Planck-Gesellschaft).


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 light and dark periods. They are classified under three groups according to the photoperiods: short-day plants, long-day plants, and day-neutral plants.


Physiology (; from Ancient Greek φύσις (physis), meaning 'nature, origin', and -λογία (-logia), meaning 'study of') is the scientific study of the functions and mechanisms which work within a living system.As a sub-discipline of biology, the focus of physiology is on how organisms, organ systems, organs, cells, and biomolecules carry out the chemical and physical functions that exist in a living system.Central to an understanding of physiological functioning is the investigation of the fundamental biophysical and biochemical phenomena, the coordinated homeostatic control mechanisms, and the continuous communication between cells.The physiologic state is the condition occurring from normal body function, while the pathological state is centered on the abnormalities that occur in animal diseases, including humans.According to the type of investigated organisms, the field can be divided into, animal physiology (including that of humans), plant physiology, cellular physiology and microbial physiology.The Nobel Prize in Physiology or Medicine is awarded to those who make significant achievements in this discipline by the Royal Swedish Academy of Sciences.

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. The current editor-in-chief is Michael R. Blatt (University of Glasgow and Laboratory of Plant Physiology and Biophysics). According to the Journal Citation Reports, the journal has a 2017 impact factor of 5.949.

Plant cuticle

A plant cuticle is a protecting film covering the epidermis of leaves, young shoots and other aerial plant organs without periderm. It consists of lipid and hydrocarbon polymers impregnated with wax, and is synthesized exclusively by the epidermal cells.


In botany and dendrology, a rhizome (, from Ancient Greek: rhízōma "mass of roots", from rhizóō "cause to strike root") is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow horizontally. The rhizome also retains the ability to allow new shoots to grow upwards.A rhizome is the main stem of the plant. A stolon is similar to a rhizome, but a stolon sprouts from an existing stem, has long internodes, and generates new shoots at the end, such as in the strawberry plant. In general, rhizomes have short internodes, send out roots from the bottom of the nodes, and generate new upward-growing shoots from the top of the nodes.

A stem tuber is a thickened part of a rhizome or stolon that has been enlarged for use as a storage organ. In general, a tuber is high in starch, e.g. the potato, which is a modified stolon. The term "tuber" is often used imprecisely and is sometimes applied to plants with rhizomes.

If a rhizome is separated each piece may be able to give rise to a new plant. The plant uses the rhizome to store starches, proteins, and other nutrients. These nutrients become useful for the plant when new shoots must be formed or when the plant dies back for the winter. This is a process known as vegetative reproduction and is used by farmers and gardeners to propagate certain plants. This also allows for lateral spread of grasses like bamboo and bunch grasses. Examples of plants that are propagated this way include hops, asparagus, ginger, irises, lily of the valley, cannas, and sympodial orchids.

Some rhizomes that are used directly in cooking include ginger, turmeric, galangal, fingerroot, and lotus.

Stored rhizomes are subject to bacterial and fungal infections, making them unsuitable for replanting and greatly diminishing stocks. However, rhizomes can also be produced artificially from tissue cultures. The ability to easily grow rhizomes from tissue cultures leads to better stocks for replanting and greater yields. The plant hormones ethylene and jasmonic acid have been found to help induce and regulate the growth of rhizomes, specifically in rhubarb. Ethylene that was applied externally was found to affect internal ethylene levels, allowing easy manipulations of ethylene concentrations. Knowledge of how to use these hormones to induce rhizome growth could help farmers and biologists producing plants grown from rhizomes more easily cultivate and grow better plants.

Some plants have rhizomes that grow above ground or that lie at the soil surface, including some Iris species, and ferns, whose spreading stems are rhizomes. Plants with underground rhizomes include gingers, bamboo, the Venus flytrap, Chinese lantern, western poison-oak, hops, and Alstroemeria, and the weeds Johnson grass, Bermuda grass, and purple nut sedge. Rhizomes generally form a single layer, but in giant horsetails, can be multi-tiered.Many rhizomes have culinary value, and some, such as zhe'ergen, are commonly consumed raw.


Sap is a fluid transported in xylem cells (vessel elements or tracheids) or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant.

Sap is distinct from latex, resin, or cell sap; it is a separate substance, separately produced, and with different components and functions.Insect honeydew is called sap, particularly when it falls from trees, but is only the remains of eaten sap and other plant parts.

Secondary growth

In botany, secondary growth is the growth that results from cell division in the cambia or lateral meristems and that causes the stems and roots to thicken, while primary growth is growth that occurs as a result of cell division at the tips of stems and roots, causing them to elongate, and gives rise to primary tissue. Secondary growth occurs in most seed plants, but monocots usually lack secondary growth. If they do have secondary growth, it differs from the typical pattern of other seed plants.

The formation of secondary vascular tissues from the cambium is a characteristic feature of dicotyledons and gymnosperms. In certain monocots, the vascular tissues are also increased after the primary growth is completed but the cambium of these plants is of a different nature. In the living Pteridophytes this feature is rare but occurs in plants like Isoetes and Botrychium.


A theca (plural thecae) refers to a sheath or a covering.


Transpiration is the process of water movement through a plant and its evaporation from aerial parts, such as leaves, stems and flowers. Water is necessary for plants but only a small amount of water taken up by the roots is used for growth and metabolism. The remaining 97–99.5% is lost by transpiration and guttation. Leaf surfaces are dotted with pores called stomata, and in most plants they are more numerous on the undersides of the foliage. The stomata are bordered by guard cells and their stomatal accessory cells (together known as stomatal complex) that open and close the pore. Transpiration occurs through the stomatal apertures, and can be thought of as a necessary "cost" associated with the opening of the stomata to allow the diffusion of carbon dioxide gas from the air for photosynthesis. Transpiration also cools plants, changes osmotic pressure of cells, and enables mass flow of mineral nutrients and water from roots to shoots. Two major factors influence the rate of water flow from the soil to the roots: the hydraulic conductivity of the soil and the magnitude of the pressure gradient through the soil. Both of these factors influence the rate of bulk flow of water moving from the roots to the stomatal pores in the leaves via the xylem.Mass flow of liquid water from the roots to the leaves is driven in part by capillary action, but primarily driven by water potential differences. If the water potential in the ambient air is lower than the water potential in the leaf airspace of the stomatal pore, water vapor will travel down the gradient and move from the leaf airspace to the atmosphere. This movement lowers the water potential in the leaf airspace and causes evaporation of liquid water from the mesophyll cell walls. This evaporation increases the tension on the water menisci in the cell walls and decrease their radius and thus the tension that is exerted on the water in the cells. Because of the cohesive properties of water, the tension travels through the leaf cells to the leaf and stem xylem where a momentary negative pressure is created as water is pulled up the xylem from the roots. In taller plants and trees, the force of gravity can only be overcome by the decrease in hydrostatic (water) pressure in the upper parts of the plants due to the diffusion of water out of stomata into the atmosphere. Water is absorbed at the roots by osmosis, and any dissolved mineral nutrients travel with it through the xylem.

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

Vascular bundle

A vascular bundle is a part of the transport system in vascular plants. The transport itself happens in vascular tissue, which exists in two forms: xylem and phloem. Both these tissues are present in a vascular bundle, which in addition will include supporting and protective tissues.

The xylem typically lies adaxial with phloem positioned abaxial. In a stem or root this means that the xylem is closer to the centre of the stem or root while the phloem is closer to the exterior. In a leaf, the adaxial surface of the leaf will usually be the upper side, with the abaxial surface the lower side. This is why aphids are typically found on the underside of a leaf rather than on the top, since the sugars manufactured by the plant are transported by the phloem, which is closer to the lower surface.

The position of vascular bundles relative to each other may vary considerably: see stele.

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: the vascular cambium and the cork cambium. All the vascular tissues within a particular plant together constitute the vascular tissue system of that plant.

The cells in vascular tissue are typically long and slender. Since the xylem and phloem function in the conduction of water, minerals, and nutrients throughout the plant, it is not surprising that their form should be similar to pipes. The individual cells of phloem are connected end-to-end, just as the sections of a pipe might be. As the plant grows, new vascular tissue differentiates in the growing tips of the plant. The new tissue is aligned with existing vascular tissue, maintaining its connection throughout the plant.

The vascular tissue in plants is arranged in long, discrete strands called vascular bundles. These bundles include both xylem and phloem, as well as supporting and protective cells. In stems and roots, the xylem typically lies closer to the interior of the stem with phloem towards the exterior of the stem. In the stems of some Asterales dicots, there may be phloem located inwardly from the xylem as well.

Between the xylem and phloem is a meristem called the vascular cambium. This tissue divides off cells that will become additional xylem and phloem. This growth increases the girth of the plant, rather than its length. As long as the vascular cambium continues to produce new cells, the plant will continue to grow more stout. In trees and other plants that develop wood, the vascular cambium allows the expansion of vascular tissue that produces woody growth. Because this growth ruptures the epidermis of the stem, woody plants also have a cork cambium that develops among the phloem. The cork cambium gives rise to thickened cork cells to protect the surface of the plant and reduce water loss. Both the production of wood and the production of cork are forms of secondary growth.

In leaves, the vascular bundles are located among the spongy mesophyll. The xylem is oriented toward the adaxial surface of the leaf (usually the upper side), and phloem is oriented toward the abaxial surface of the leaf. This is why aphids are typically found on the undersides of the leaves rather than on the top, since the phloem transports sugars manufactured by the plant and they are closer to the lower surface.

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