|This article needs additional citations for verification. (February 2010)|
||This article duplicates, in whole or part, the scope of other articles, specifically, Soil#Nutrients.|
Plant nutrition is the study of the chemical elements and compounds that are necessary for plant growth, and also of their external supply and internal metabolism. In 1972, E. Epstein defined two criteria for an element to be essential for plant growth:
- in its absence the plant is unable to complete a normal life cycle; or
- that the element is part of some essential plant constituent or metabolite.
This is in accordance with Liebig's law of the minimum. There are 14 essential plant nutrients. Carbon and oxygen are absorbed from the air, while other nutrients including water are obtained from the soil. Plants must obtain the following mineral nutrients from the growing media:
- the primary macronutrients: nitrogen (N), phosphorus (P), potassium (K)
- the three secondary macronutrients: calcium (Ca), sulphur (S), magnesium (Mg)
- the micronutrients/trace minerals: boron (B), chlorine (Cl), manganese (Mn), iron (Fe), zinc (Zn), copper (Cu), molybdenum (Mo), nickel (Ni)
The macronutrients are consumed in larger quantities and are present in plant tissue in quantities from 0.2% to 4.0% (on a dry matter weight basis). Micro nutrients are present in plant tissue in quantities measured in parts per million, ranging from 5 to 200 ppm, or less than 0.02% dry weight.
Most soil conditions across the world can provide plants with adequate nutrition and do not require fertilizer for a complete life cycle. However, humans can artificially modify soil through the addition of fertilizer to promote vigorous growth and increase yield. The plants are able to obtain their required nutrients from the fertilizer added to the soil. A colloidal carbonaceous residue, known as humus, can serve as a nutrient reservoir. Even with adequate water and sunshine, nutrient deficiency can limit growth.
Nutrient uptake from the soil is achieved by cation exchange, where root hairs pump hydrogen ions (H+) into the soil through proton pumps. These hydrogen ions displace cations attached to negatively charged soil particles so that the cations are available for uptake by the root.
Plant nutrition is a difficult subject to understand completely, partly because of the variation between different plants and even between different species or individuals of a given clone. An element present at a low level may cause deficiency symptoms, while the same element at a higher level may cause toxicity. Further, deficiency of one element may present as symptoms of toxicity from another element. An abundance of one nutrient may cause a deficiency of another nutrient. For example, lower availability of a given nutrient such as SO24− can affect the uptake of another nutrient, such as NO3–. As another example, K+ uptake can be influenced by the amount of NH4+ available.
The root, especially the root hair, is the most essential organ for the uptake of nutrients. The structure and architecture of the root can alter the rate of nutrient uptake. Nutrient ions are transported to the center of the root, the stele in order for the nutrients to reach the conducting tissues, xylem and phloem. The Casparian strip, a cell wall outside of the stele but within the root, prevents passive flow of water and nutrients, helping to regulate the uptake of nutrients and water. Xylem moves water and inorganic molecules within the plant and phloem accounts for organic molecule transportation. Water potential plays a key role in a plants nutrient uptake. If the water potential is more negative within the plant than the surrounding soils, the nutrients will move from the region of higher solute concentration--in the soil--to the area of lower solute concentration: in the plant.
There are three fundamental ways plants uptake nutrients through the root: 1.) simple diffusion, occurs when a nonpolar molecule, such as O2, CO2, and NH3 follows a concentration gradient, moving passively through the cell lipid bilayer membrane without the use of transport proteins. 2.) facilitated diffusion, is the rapid movement of solutes or ions following a concentration gradient, facilitated by transport proteins. 3.) Active transport, is the uptake by cells of ions or molecules against a concentration gradient; this requires an energy source, usually ATP, to power molecular pumps that move the ions or molecules through the membrane.
- Nutrients are moved inside a plant to where they are most needed. For example, a plant will try to supply more nutrients to its younger leaves than to its older ones. When nutrients are mobile, symptoms of any deficiency become apparent first on the older leaves. However, not all nutrients are equally mobile. Nitrogen, phosphorus, and potassium are mobile nutrients, while the others have varying degrees of mobility. When a less mobile nutrient is deficient, the younger leaves suffer because the nutrient does not move up to them but stays in the older leaves. This phenomenon is helpful in determining which nutrients a plant may be lacking.
Many plants engage in symbiosis with microorganisms. Two important types of these relationship are 1.) with bacteria such as rhizobia, that carry out biological nitrogen fixation, in which atmospheric nitrogen (N2) is converted into ammonium (NH4); and 2.) with mycorrhizal fungi, which through their association with the plant roots help to create a larger effective root surface area. Both of these mutualistic relationships enhance nutrient uptake.
Though nitrogen is plentiful in the Earth's atmosphere, relatively few plants harbor nitrogen fixing bacteria, so most plants rely on nitrogen compounds present in the soil to support their growth. These can be supplied by mineralization of soil organic matter or added plant residues, nitrogen fixing bacteria, animal waste, or through the application of fertilizers.
Hydroponics, is a method for growing plants in a water-nutrient solution without the use of nutrient-rich soil. It allows researchers and home gardeners to grow their plants in a controlled environment. The most common solution, is the Hoagland solution, developed by D. R. Hoagland in 1933, the solution consists of all the essential nutrients in the correct proportions necessary for most plant growth. An aerator is used to prevent an anoxic event or hypoxia. Hypoxia can affect nutrient uptake of a plant because without oxygen present, respiration becomes inhibited within the root cells. The Nutrient film technique is a variation of hydroponic technique. The roots are not fully submerged, which allows for adequate aeration of the roots, while a "film" thin layer of nutrient rich water is pumped through the system to provide nutrients and water to the plant.
- 1 Processes
- 2 Functions of nutrients
- 2.1 Macronutrients (derived from air and water)
- 2.2 Macronutrients (primary)
- 2.3 Macronutrients (secondary and tertiary)
- 2.4 Micro-nutrients
- 3 See also
- 4 References
- 5 External links
Plants take up essential elements from the soil through their roots and from the air (mainly consisting of nitrogen and oxygen) through their leaves. Nutrient uptake in the soil is achieved by cation exchange, wherein root hairs pump hydrogen ions (H+) into the soil through proton pumps. These hydrogen ions displace cations attached to negatively charged soil particles so that the cations are available for uptake by the root. In the leaves, stomata open to take in carbon dioxide and expel oxygen. The carbon dioxide molecules are used as the carbon source in photosynthesis.
Functions of nutrients
Although nitrogen is plentiful in the Earth's atmosphere, relatively few plants engage in nitrogen fixation (conversion of atmospheric nitrogen to a biologically useful form). Most plants therefore require nitrogen compounds to be present in the soil in which they grow.
Plant nutrition is a difficult subject to understand completely, partially because of the variation between different plants and even between different species or individuals of a given clone. Elements present at low levels may cause deficiency symptoms, and toxicity is possible at levels that are too high. Furthermore, deficiency of one element may present as symptoms of toxicity from another element, and vice versa.
Carbon and oxygen are absorbed from the air, while other nutrients are absorbed from the soil. Green plants obtain their carbohydrate supply from the carbon dioxide in the air by the process of photosynthesis. Each of these nutrients is used in a different place for a different essential function.
Macronutrients (derived from air and water)
Carbon forms the backbone of many plants biomolecules, including starches and cellulose. Carbon is fixed through photosynthesis from the carbon dioxide in the air and is a part of the carbohydrates that store energy in the plant.
Hydrogen also is necessary for building sugars and building the plant. It is obtained almost entirely from water. Hydrogen ions are imperative for a proton gradient to help drive the electron transport chain in photosynthesis and for respiration.
Oxygen by itself or in the molecules of H2O or CO2 are necessary for plant cellular respiration. Cellular respiration is the process of generating energy-rich adenosine triphosphate (ATP) via the consumption of sugars made in photosynthesis. Plants produce oxygen gas during photosynthesis to produce glucose but then require oxygen to undergo aerobic cellular respiration and break down this glucose and produce ATP.
Phosphorus is important in plant bioenergetics. As a component of ATP, phosphorus is needed for the conversion of light energy to chemical energy (ATP) during photosynthesis. Phosphorus can also be used to modify the activity of various enzymes by phosphorylation, and can be used for cell signaling. Since ATP can be used for the biosynthesis of many plant biomolecules, phosphorus is important for plant growth and flower/seed formation. Phosphate esters make up DNA, RNA, and phospholipids. Most common in the form of polyprotic phosphoric acid (H3PO4) in soil, but it is taken up most readily in the form of H2PO4. Phosphorus is limited in most soils because it is released very slowly from insoluble phosphates. Under most environmental conditions it is the limiting element because of its small concentration in soil and high demand by plants and microorganisms. Plants can increase phosphorus uptake by a mutualism with mycorrhiza. A Phosphorus deficiency in plants is characterized by an intense green coloration in leaves. If the plant is experiencing high phosphorus deficiencies the leaves may become denatured and show signs of necrosis. Occasionally the leaves may appear purple from an accumulation of anthocyanin. Because phosphorus is a mobile nutrient, older leaves will show the first signs of deficiency.
It is useful to apply a high phosphorus content fertilizer, such as bone meal, to perennials to help with successful root formation.
Potassium regulates the opening and closing of the stomata by a potassium ion pump. Since stomata are important in water regulation, potassium reduces water loss from the leaves and increases drought tolerance. Potassium deficiency may cause necrosis or interveinal chlorosis. K+ is highly mobile and can aid in balancing the anion charges within the plant. Potassium helps in fruit colouration, shape and also increases its brix. Hence, quality fruits are produced in Potassium rich soils. It also has high solubility in water and leaches out of rocky or sandy soils. This water solubility can result in potassium deficiency. Potassium serves as an activator of enzymes used in photosynthesis and respiration Potassium is used to build cellulose and aids in photosynthesis by the formation of a chlorophyll precursor. Potassium deficiency may result in higher risk of pathogens, wilting, chlorosis, brown spotting, and higher chances of damage from frost and heat.
Nitrogen is an essential component of all proteins. Nitrogen deficiency most often results in stunted growth, slow growth, and chlorosis. Nitrogen deficient plants will also exhibit a purple appearance on the stems, petioles and underside of leaves from an accumulation of anthocyanin pigments. Most of the nitrogen taken up by plants is from the soil in the forms of NO3–, although in acid environments such as boreal forests where nitrification is less likely to occur, ammonium NH4+ is more likely to be the dominating source of nitrogen. Amino acids and proteins can only be built from NH4+ so NO3– must be reduced. Under many agricultural settings, nitrogen is the limiting nutrient of high growth. Some plants require more nitrogen than others, such as corn (Zea mays). Because nitrogen is mobile, the older leaves exhibit chlorosis and necrosis earlier than the younger leaves. Soluble forms of nitrogen are transported as amines and amides.
Macronutrients (secondary and tertiary)
Sulphur is a structural component of some amino acids and vitamins, and is essential in the manufacturing of chloroplasts. Sulphur is also found in the Iron Sulphur complexes of the electron transport chains in photosynthesis. It is immobile and deficiency therefore affects younger tissues first. Symptoms of deficiency include yellowing of leaves and stunted growth.
Calcium regulates transport of other nutrients into the plant and is also involved in the activation of certain plant enzymes. Calcium deficiency results in stunting. This nutrient is involved in photosynthesis and plant structure. Blossom end rot is also a result of inadequate calcium.
Magnesium is an important part of chlorophyll, a critical plant pigment important in photosynthesis. It is important in the production of ATP through its role as an enzyme cofactor. Magnesium deficiency can result in interveinal chlorosis.
In plants, silicon strengthens cell walls, improving plant strength, health, and productivity. Other benefits of silicon to plants include improved drought and frost resistance, decreased lodging potential and boosting the plant's natural pest and disease fighting systems. Silicon has also been shown to improve plant vigor and physiology by improving root mass and density, and increasing above ground plant biomass and crop yields. Although not considered an essential element for plant growth and development (except for specific plant species - sugarcane and members of the horsetail family), silicon is considered a beneficial element in many countries throughout the world due to its many benefits to numerous plant species when under abiotic or biotic stresses. Silicon is currently under consideration by the Association of American Plant Food Control Officials (AAPFCO) for elevation to the status of a "plant beneficial substance".
Silicon is the second most abundant element in earth's crust. Higher plants differ characteristically in their capacity to take up silicon. Depending on their SiO2 content they can be divided into three major groups:
- Wetland graminae-wetland rice, horsetail (10–15%)
- Dryland graminae-sugar cane, most of the cereal species and few dicotyledons species (1–3%)
- Most of dicotyledons especially legumes (<0.5%)
- The long distance transport of Si in plants is confined to the xylem. Its distribution within the shoot organ is therefore determined by transpiration rate in the organs
- The epidermal cell walls are impregnated with a film layer of silicon and effective barrier against water loss, cuticular transpiration rate in the organs.
Si can stimulate growth and yield by several indirect actions. These include decreasing mutual shading by improving leaf erectness, decreasing susceptibility to lodging, preventing Mn and Fe toxicity.
Some elements are directly involved in plant metabolism (Arnon and Stout, 1939). However, this principle does not account for the so-called beneficial elements, whose presence, while not required, has clear positive effects on plant growth. Mineral elements that either stimulate growth but are not essential, or that are essential only for certain plant species, or under given conditions, are usually defined as beneficial elements.
Iron is necessary for photosynthesis and is present as an enzyme cofactor in plants. Iron deficiency can result in interveinal chlorosis and necrosis. Iron is not a structural part of chlorophyll but very much essential for its synthesis. Copper deficiency can be responsible for promoting an iron deficiency.
Molybdenum is a cofactor to enzymes important in building amino acids. Involved in Nitrogen metabolism. Mo is part of Nitrate reductase enzyme.
Boron is important for binding of pectins in the RGII region of the primary cell wall, secondary roles may be in sugar transport, cell division, and synthesizing certain enzymes. Boron deficiency causes necrosis in young leaves and stunting.
Copper is important for photosynthesis. Symptoms for copper deficiency include chlorosis. Involved in many enzyme processes. Necessary for proper photosythesis. Involved in the manufacture of lignin (cell walls). Involved in grain production. It is also hard to find in some conditions.
- Essential for C4 plants rather C3
- Substitution of K by Na: Plants can be classified into four groups:
- Group A—a high proportion of K can be replaced by Na and stimulate the growth, which cannot be achieved by the application of K
- Group B—specific growth responses to Na are observed but they are much less distinct
- Group C—Only minor substitution is possible and Na has no effect
- Group D—No substitution is occurred
- Stimulate the growth—increase leaf area, stomata, improve the water balance
- Na functions in metabolism
- C4 metabolism
- Impair the conversion of pyruvate to phosphoenol-pyruva
- Reduce the photosystem II activity and ultrastructural changes in mesophyll chloroplast
- Replacing K functions
- Internal osmoticum
- Stomatal function
- Counteraction in long distance transport
- Enzyme activation
- Improves the crop quality e.g. improve the taste of carrots by increasing sucrose
Zinc is required in a large number of enzymes and plays an essential role in DNA transcription. A typical symptom of zinc deficiency is the stunted growth of leaves, commonly known as "little leaf" and is caused by the oxidative degradation of the growth hormone auxin.
In higher plants, Nickel is absorbed by plants in the form of Ni2+ ion. Nickel is essential for activation of urease, an enzyme involved with nitrogen metabolism that is required to process urea. Without Nickel, toxic levels of urea accumulate, leading to the formation of necrotic lesions. In lower plants, Nickel activates several enzymes involved in a variety of processes, and can substitute for Zinc and Iron as a cofactor in some enzymes.
Cobalt has proven to be beneficial to at least some plants, but is essential in others, such as legumes where it is required for nitrogen fixation for the symbiotic relationship it has with nitrogen-fixing bacteria. Vanadium may be required by some plants, but at very low concentrations. It may also be substituting for molybdenum. Selenium and sodium may also be beneficial. Sodium can replace potassium's regulation of stomatal opening and closing.
- The requirement of Co for N2 fixation in legumes and non-legumes have been documented clearly
- Protein synthesis of Rhizobium is impaired due to Co deficiency
- It is still not clear whether Co has direct effect on higher plant
- Tea has a high tolerance for Al toxicity and the growth is stimulated by Al application. The possible reason is the prevention of Cu, Mn or P toxicity effects.
- There have been reports that Al may serve as fungicide against certain types of root rot.
- Emanuel Epstein. Mineral Nutrition of Plants: Principles and Perspectives.
- Allen V. Barker; D. J. Pilbeam (2007). Handbook of plant nutrition. CRC Press. pp. 4–. ISBN 978-0-8247-5904-9. Retrieved 17 August 2010.
- http://aesl.ces.uga.edu/publications/plant/Nutrient.htm Retrieved Jan. 2010
- Norman P. A. Huner; William Hopkins. "3 & 4". Introduction to Plant Physiology 4th Edition. John Wiley & Sons, Inc. ISBN 978-0-470-24766-2.
- Pages 68 and 69 Taiz and Zeiger Plant Physiology 3rd Edition 2002 ISBN 0-87893-823-0
- Lowenfels, Lewis, Jeff, Wayne (2011). Teaming with microbes. pp. 49, 110. ISBN 978-1-60469-113-9.
- University of Zurich (2011). Blossom end rot: Transport protein identified. http://phys.org/news/2011-11-blossom-protein.html
- (2012). New Light Shined on Photosynthesis. http://www.newswise.com/articles/new-light-shined-on-photosynthesis University of Arizona
- "Silicon nutrition in plants". Plant Health Care,Inc.: 1. 12 December 2000. Retrieved 1 July 2011.
- Prakash, Dr. N. B. (2007). "Evaluation of the calcium silicate as a source of silicon in aerobic and wet rice". University of Agricultural Science Bangalore. p. 1.
- AgriPower. "A Review of Silicon and Its Benefits for Plants". pp. 38–41. Retrieved 19 July 2011.
- Feng Ma, Jian; Yamaji, Naoki (12 July 2006). "Silicon uptake and accumulation in higher plants". Trend in Plant Science. Abiotic stress series 11 (8): 1. Retrieved 1 July 2011.
- Feng Ma, Jian; Yamaji, Naoki (12 July 2006). "Silicon uptake and accumulation in higher plants". Trend in Plant Science. Abiotic stress series 11 (8): 4–5. Retrieved 1 July 2011.
- "AAPFCO Board of Directors 2006 Mid-Year Meeting". Association of American Plant Food Control Officials. Retrieved 18 July 2011.
- Miranda, Stephen R.; Barker, Bruce (August 4, 2009). "Silicon: Summary of Extraction Methods". Harsco Minerals. Retrieved 18 July 2011.
- (2012). "Nutrient and toxin all at once: How plants absorb the perfect quantity of minerals". http://esciencenews.com/articles/2012/04/13/nutrient.and.toxin.all.once.how.plants.absorb.perfect.quantity.minerals Ruhr-Universität
- (2012).New Light Shined on Photosynthesis. http://www.newswise.com/articles/new-light-shined-on-photosynthesis Arizona State University.
- Journal of Plant Nutrition
- International Fertilizer Industry Association
- Principles of Plant Nutrition