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 in 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 either be supplied by decaying matter, nitrogen fixing bacteria, animal waste, or through the agricultural application of purpose-made 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.
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