Hormone
A hormone (from the Greek participle ὁρμῶν, "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs or tissues by complex biological processes to regulate physiology and behavior.[1] Hormones are required for the correct development of animals, plants and fungi. Due to the broad definition of a hormone (as a signaling molecule that exerts its effects far from its site of production), numerous kinds of molecules can be classified as hormones. Among the substances that can be considered hormones, are eicosanoids (e.g. prostaglandins and thromboxanes), steroids (e.g. oestrogen and brassinosteroid), amino acid derivatives (e.g. epinephrine and auxin), protein or peptides (e.g. insulin and CLE peptides), and gases (e.g. ethylene and nitric oxide).
For other uses, see Hormone (disambiguation).
Hormones are used to communicate between organs and tissues. In vertebrates, hormones are responsible for regulating a wide range of processes including both physiological processes and behavioral activities such as digestion, metabolism, respiration, sensory perception, sleep, excretion, lactation, stress induction, growth and development, movement, reproduction, and mood manipulation.[2][3][4] In plants, hormones modulate almost all aspects of development, from germination to senescence.[5]
Hormones affect distant cells by binding to specific receptor proteins in the target cell, resulting in a change in cell function. When a hormone binds to the receptor, it results in the activation of a signal transduction pathway that typically activates gene transcription, resulting in increased expression of target proteins. Hormones can also act in non-genomic pathways that synergize with genomic effects.[6] Water-soluble hormones (such as peptides and amines) generally act on the surface of target cells via second messengers. Lipid soluble hormones, (such as steroids) generally pass through the plasma membranes of target cells (both cytoplasmic and nuclear) to act within their nuclei. Brassinosteroids, a type of polyhydroxysteroids, are a sixth class of plant hormones and may be useful as an anticancer drug for endocrine-responsive tumors to cause apoptosis and limit plant growth. Despite being lipid soluble, they nevertheless attach to their receptor at the cell surface.[7]
In vertebrates, endocrine glands are specialized organs that secrete hormones into the endocrine signaling system. Hormone secretion occurs in response to specific biochemical signals and is often subject to negative feedback regulation. For instance, high blood sugar (serum glucose concentration) promotes insulin synthesis. Insulin then acts to reduce glucose levels and maintain homeostasis, leading to reduced insulin levels. Upon secretion, water-soluble hormones are readily transported through the circulatory system. Lipid-soluble hormones must bond to carrier plasma glycoproteins (e.g., thyroxine-binding globulin (TBG)) to form ligand-protein complexes. Some hormones, such as insulin and growth hormones, can be released into the bloodstream already fully active. Other hormones, called prohormones, must be activated in certain cells through a series of steps that are usually tightly controlled.[8] The endocrine system secretes hormones directly into the bloodstream, typically via fenestrated capillaries, whereas the exocrine system secretes its hormones indirectly using ducts. Hormones with paracrine function diffuse through the interstitial spaces to nearby target tissue.
Plants lack specialized organs for the secretion of hormones, although there is spatial distribution of hormone production. For example, the hormone auxin is produced mainly at the tips of young leaves and in the shoot apical meristem. The lack of specialised glands means that the main site of hormone production can change throughout the life of a plant, and the site of production is dependent on the plant's age and environment.[9]
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Hormones have the following effects on the body:[31]
A hormone may also regulate the production and release of other hormones. Hormone signals control the internal environment of the body through homeostasis.
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The rate of hormone biosynthesis and secretion is often regulated by a homeostatic negative feedback control mechanism. Such a mechanism depends on factors that influence the metabolism and excretion of hormones. Thus, higher hormone concentration alone cannot trigger the negative feedback mechanism. Negative feedback must be triggered by overproduction of an "effect" of the hormone.[32][33]
Hormone secretion can be stimulated and inhibited by:
One special group of hormones is the tropic hormones that stimulate the hormone production of other endocrine glands. For example, thyroid-stimulating hormone (TSH) causes growth and increased activity of another endocrine gland, the thyroid, which increases output of thyroid hormones.[34]
To release active hormones quickly into the circulation, hormone biosynthetic cells may produce and store biologically inactive hormones in the form of pre- or prohormones. These can then be quickly converted into their active hormone form in response to a particular stimulus.[34]
Eicosanoids are considered to act as local hormones. They are considered to be "local" because they possess specific effects on target cells close to their site of formation. They also have a rapid degradation cycle, making sure they do not reach distant sites within the body.[35]
Hormones are also regulated by receptor agonists. Hormones are ligands, which are any kinds of molecules that produce a signal by binding to a receptor site on a protein. Hormone effects can be inhibited, thus regulated, by competing ligands that bind to the same target receptor as the hormone in question. When a competing ligand is bound to the receptor site, the hormone is unable to bind to that site and is unable to elicit a response from the target cell. These competing ligands are called antagonists of the hormone.[36]
At the neurological level, behavior can be inferred based on hormone concentration, which in turn are influenced by hormone-release patterns; the numbers and locations of hormone receptors; and the efficiency of hormone receptors for those involved in gene transcription. Hormone concentration does not incite behavior, as that would undermine other external stimuli; however, it influences the system by increasing the probability of a certain event to occur.[39]
Not only can hormones influence behavior, but also behavior and the environment can influence hormone concentration.[40] Thus, a feedback loop is formed, meaning behavior can affect hormone concentration, which in turn can affect behavior, which in turn can affect hormone concentration, and so on.[41] For example, hormone-behavior feedback loops are essential in providing constancy to episodic hormone secretion, as the behaviors affected by episodically secreted hormones directly prevent the continuous release of sad hormones.[42]
Three broad stages of reasoning may be used to determine if a specific hormone-behavior interaction is present within a system:
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Though colloquially oftentimes used interchangeably, there are various clear distinctions between hormones and neurotransmitters:[43][44][36]
Neurohormones are a type of hormone that share a commonality with neurotransmitters.[47] They are produced by endocrine cells that receive input from neurons, or neuroendocrine cells.[47] Both classic hormones and neurohormones are secreted by endocrine tissue; however, neurohormones are the result of a combination between endocrine reflexes and neural reflexes, creating a neuroendocrine pathway.[36] While endocrine pathways produce chemical signals in the form of hormones, the neuroendocrine pathway involves the electrical signals of neurons.[36] In this pathway, the result of the electrical signal produced by a neuron is the release of a chemical, which is the neurohormone.[36] Finally, like a classic hormone, the neurohormone is released into the bloodstream to reach its target.[36]
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