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Plant root exudates

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Plant root exudates

Plant root exudates are fluids emitted through the roots of plants. These secretions influence the rhizosphere around the roots to inhibit harmful microbes and promote the growth of self and kin plants.

Plant root systems can grow to be complex due to a variety of species and microorganisms existing in a common soil. Plants have adapted to respond to the soil conditions and presence of microbes through various mechanisms, one of which is the secretion of root exudates. This secretion allows plants to largely influence the rhizosphere as well as the organisms that exist within it. The contents of exudates and the amount of substance released is reliant on multiple factors, including the root system architecture, presence of harmful microbes, and metal toxicity. The species of the plant as well as its developmental stage can also influence the chemical mixture that is released through exudates. The contents may include ions, carbon-based compounds, amino acids, sterols, and many other chemical compounds. At sufficient concentrations, exudates are capable of mediating[clarification needed] both positive and negative plant-plant and plant-microbe interactions.

The physiological mechanism by which exudates are released is not entirely understood and varies depending on the stimulus as well as the contents of the secreted exudate. Various types of root cells have been suggested to sense microbes or compounds in the soil and secrete exudates accordingly. One example of root exudation occurs when plants sense elicitors and prime[clarification needed] for a stress or defense response.[clarification needed] It is believed that elicitors, such as methyl jasmonate and salicylic acid, are sensed by receptors on root cap cells, often referred to as border cells. This induces a change in gene regulation, up-regulating specific defense or stress-response genes. This differential gene expression results in metabolic changes that ultimately result in the biosynthesis of primary and secondary metabolites. These metabolites exit cells in the form of exudates through transporters that vary depending on the chemical structure of the metabolites. The exudate secretion is then able to elicit a defense response against harmful microbes within the soil.

The rhizosphere is the thin area of soil immediately surrounding the root system. It is a densely populated area in which the roots compete with invading root systems of neighboring plant species for space, water, and mineral nutrients as well as form positive and negative relationships with soil-borne microorganisms such as bacteria, fungi and insects. Quantifying how much photosynthetically fixed carbon is transferred to soil via plant root exudates is difficult, but 5% can be considered a rough estimate. Root exudates are seen as key mediators in the interaction between plants and soil microbiota.

Root exudates contain a wide variety of molecules released by the plant into the soil. They act as a signaling messenger that allows for communication between soil microbes and plant roots.[citation needed] Exudates influence several factors within the soil such as nutrient availability, soil pH, and recruitment of bacteria and fungi. All of these impact the relationships that plants have with each other as well as soilborne microorganisms. The most notable positive relationship is that of roots and mycorrhizae. It is estimated that 80-90% of plants are colonized by mycorrhizae in nature. Mycorrhizae are known to promote plant growth and increase water use efficiency. Plants establish these mutualistic relationships with bacteria and fungi by modulating the composition of the root exudates. While positive relationships like this do exist, it is worth noting that most microbes have incompatible interactions with plants. One of the main forms of negative relationships in the rhizosphere is allelopathy. This is the act of releasing phytotoxins into the rhizosphere that can influence neighboring plant's growth, respiration, photosynthesis, metabolism, and water and nutrient uptake. Allelochemicals released by the roots do this by inducing changes in cell structures, inhibiting cell division and elongation, destabilizing the antioxidant system, and increasing membrane permeability.

The plant family (Asteraceae, Brassicaceae, Fabaceae, and Poaceae) is the most important source of variation in exudation rates and microbial community structure between plant species. Root symbiotic associations impacts the rate of sugar exudation in the rhizosphere. Root exudation impacts microbial activities as well as the diversity of active microbiota involved in root exudate assimilation. Root exudates play a major role in root-soil contact, the exact purpose of the exudates and the reactions they cause are still poorly understood.

Plants have developed various advantageous mechanisms to manipulate their habitats. This is important as a plants’ habitat is crucial towards their growth as it dictates energy, water intake, nutrient intake and others [1]. Thus, a mechanism known as exudation that has been used by plants to possibly manipulate its surroundings, has been found to be useful although it is not fully understood how plants utilize it. Nor is it understood if the process of exudations is truly advantageous or how it is controlled by plants. An example of this would be the maize species which is grown as an agricultural staple and thus is located in close proximity to other species of plants. The maize plant releases exudates to deter herbivore attacks from pest by reducing its leaf nutrient value and as well repressing its size. While a defensive mechanism this action can be problematic for farmers because repression of growth affects their bounties. A number of proposals to explain this mechanism have been offered however, they are just proposals and have not been fully developed and tested to assert their claims. One such claim is that root exudates are beneficial for defense. While another claims that exudates can also recognize who is related to the plant and who is a stranger which promotes friendly competition. Another claim asserts that plants are able to possibly adjust their source-sink allocation of resources and the process of exudation which promote positive effects for the plant's growth[2].

Primary metabolites that are released into the soil by plants consist of: amino acids, organic acids and sugars. These primary metabolites are thought to be primarily released through the root tip when the rhizosphere is negatively affected by stressors such as being nutrient poor. This environmental sense of surroundings allows the plant to dictate when these metabolites should be released. The mechanism described for this process is illustrated by facilitated diffusion from the root tip, this process requires the possible adjustment of the source sink conserves and this creates a pressure driven mechanism through the phloem. Traveling through the simplistic pathway is the most common method as they can travel freely however while nearing their journey, they have to pass through a plasma membrane and to do this they need a transmembrane protein to complete the trip. “The phloem unloads the primary metabolites through the plasmodesmata using both facilitated diffusion and pressure flow mechanics to push release at the root tip”.

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