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In genetics , the phenotype (from Ancient Greek φαίνω ( phaínō )  'to appear, show' and τύπος ( túpos )  'mark, type') is the set of observable characteristics or traits of an organism . The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological properties, its behavior , and the products of behavior. An organism's phenotype results from two basic factors: the expression of an organism's genetic code (its genotype ) and the influence of environmental factors. Both factors may interact, further affecting the phenotype. When two or more clearly different phenotypes exist in the same population of a species, the species is called polymorphic . A well-documented example of polymorphism is Labrador Retriever coloring ; while the coat color depends on many genes, it is clearly seen in the environment as yellow, black, and brown. Richard Dawkins in 1978 and then again in his 1982 book The Extended Phenotype suggested that one can regard bird nests and other built structures such as caddisfly larva cases and beaver dams as "extended phenotypes".

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49-544: Maca is an edible plant endemic to the Andes. MACA or maca can mean: Maca Lepidium peruvianum Lepidium meyenii , known as maca or Peruvian ginseng , is an edible herbaceous biennial plant of the family Brassicaceae that is native to South America in the high Andes mountains of Peru and Bolivia . It was rediscovered for commercial purposes at the Meseta de Bombón plateau close to Lake Junin in

98-423: A U.S.-based brewery called Andean Brewing Company became the first company to produce and commercialize beer made from maca under the brand KUKA Beer. From the black morphotype, a liquor is produced. Also, the leaves are edible or may serve as animal fodder. They can be prepared raw in salads or cooked much like L. sativum and L. campestre , which are closely related genetically. The prominent product for export

147-484: A bird feeds a brood parasite such as a cuckoo , it is unwittingly extending its phenotype; and when genes in an orchid affect orchid bee behavior to increase pollination, or when genes in a peacock affect the copulatory decisions of peahens, again, the phenotype is being extended. Genes are, in Dawkins's view, selected by their phenotypic effects. Other biologists broadly agree that the extended phenotype concept

196-444: A central raceme , and are followed by 4–5 mm (0.16–0.20 in) siliculate fruits, each containing two small 2.0–2.5 mm (0.079–0.098 in) reddish-gray ovoid seeds . Seeds are the maca's only means of reproduction. Maca reproduces mainly through self-pollination and is an autogamous species. The genome consists of 64 chromosomes. Maca is a short-day plant from experiments with different day lengths. Some sources consider

245-631: A cultivated hectare may reach an estimated 15 tons in fresh hypocotyls, resulting in around 5 tons of dried material. According to the Ministry of Agriculture of Peru , however, average maca yields for 2005 were only 7 t/ha, with a great variation between different sites. Although maca has been cultivated outside the Andes, it is unclear whether it develops the same active constituents or potency outside of its natural habitat. Hypocotyls grown from Peruvian seeds form with difficulty at low elevations, in greenhouses, or warm climates. The average composition, on

294-671: A dry matter basis, is 60–75% carbohydrates (primarily as polysaccharides ), 10–14% protein , 8.5% dietary fiber , and 2.2% fats . Maca contains glucotropaeolin , m -methoxyglucotropaeolin, benzyl glucosinolates , polyphenols , (1 R ,3 S )-1-methyl-1,2,3,4-tetrahydro-β-carboline-3-carboxylic acid (MTCA), and p -methoxybenzyl isothiocyanate . Alkamides are also present in maca. Maca contains several N-benzylamides referred to as macamides that are structurally related to anandamide . No pharmacokinetic data have been published for maca. The presence of (1 R ,3 S )-1-methyl-1,2,3,4-tetrahydro-β-carboline-3-carboxylic acid (MTCA) in

343-518: A gene has on its surroundings, including other organisms, as an extended phenotype, arguing that "An animal's behavior tends to maximize the survival of the genes 'for' that behavior, whether or not those genes happen to be in the body of the particular animal performing it." For instance, an organism such as a beaver modifies its environment by building a beaver dam ; this can be considered an expression of its genes , just as its incisor teeth are—which it uses to modify its environment. Similarly, when

392-529: A large part of the Human Genome Project . Phenomics has applications in agriculture. For instance, genomic variations such as drought and heat resistance can be identified through phenomics to create more durable GMOs. Phenomics may be a stepping stone towards personalized medicine , particularly drug therapy . Once the phenomic database has acquired enough data, a person's phenomic information can be used to select specific drugs tailored to

441-537: A multidimensional search space with several neurobiological levels, spanning the proteome, cellular systems (e.g., signaling pathways), neural systems and cognitive and behavioural phenotypes." Plant biologists have started to explore the phenome in the study of plant physiology. In 2009, a research team demonstrated the feasibility of identifying genotype–phenotype associations using electronic health records (EHRs) linked to DNA biobanks . They called this method phenome-wide association study (PheWAS). Inspired by

490-453: A particular enzyme is expressed at high levels, the organism may produce more of that enzyme and exhibit a particular trait as a result. On the other hand, if the gene is expressed at low levels, the organism may produce less of the enzyme and exhibit a different trait. Gene expression is regulated at various levels and thus each level can affect certain phenotypes, including transcriptional and post-transcriptional regulation. Changes in

539-427: Is both sweet and slightly bitter. The natural environment of the maca is at 11–12ºS latitude and at an elevation of 3,800–4,400 m (12,500–14,400 ft) above sea level. At this elevation, temperatures of the growing season vary from −2 to 13 °C (28 to 55 °F) in monthly mean minimum or maximum, respectively. Temperatures can decline, however, as low as −10 °C (14 °F) and frosts are common. Of

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588-463: Is found growing in two different habitats in Sweden. One habitat is rocky, sea-side cliffs , where the plants are bushy with broad leaves and expanded inflorescences ; the other is among sand dunes where the plants grow prostrate with narrow leaves and compact inflorescences. These habitats alternate along the coast of Sweden and the habitat that the seeds of Hieracium umbellatum land in, determine

637-421: Is maca flour , a baking flour ground from the hard, dried roots. It is called harina de maca . Maca flour (powder) is a relatively inexpensive bulk commodity, like wheat flour or potato flour. The supplement industry uses both dry roots and maca flour for different types of processing and concentrated extracts. Another common form is maca, processed by gelatinization . This extrusion process separates and removes

686-403: Is mainly grown for the consumption of its roots. The majority of harvested maca is dried. In this form, the hypocotyls can be stored for several years. In Peru, maca is prepared and consumed in various ways, although traditionally, it is always cooked. The freshly harvested hypocotyl may be roasted in a pit (called huatia ) and is considered a delicacy. Fresh roots usually are available only in

735-470: Is not necessary. The climate is not suitable for most weeds or pests. Nearly all maca cultivation in Peru is carried out organically; maca is seldom attacked. Maca is sometimes interplanted with potatoes, as it is known to maca farmers that the plant naturally repels most root crop pests. The harvest is done manually, with the leaves left in the field as livestock feed or organic fertilizer . The yield for

784-417: Is problematic. A proposed definition for both terms as the "physical totality of all traits of an organism or of one of its subsystems" was put forth by Mahner and Kary in 1997, who argue that although scientists tend to intuitively use these and related terms in a manner that does not impede research, the terms are not well defined and usage of the terms is not consistent. Some usages of the term suggest that

833-469: Is related, but it also resembles a parsnip . The green, fragrant tops are short and lie along the ground. The thin, frilly leaves sprout in a rosette at the soil surface, not growing more than 12–20 cm (4.7–7.9 in) in height. The leaves show a dimorphism according to the reproductive stage. They are more prominent in the vegetative phase and continuously renew from the center as the outer leaves die. The off-white, self-fertile flowers are borne on

882-478: Is relevant, but consider that its role is largely explanatory, rather than assisting in the design of experimental tests. Phenotypes are determined by an interaction of genes and the environment, but the mechanism for each gene and phenotype is different. For instance, an albino phenotype may be caused by a mutation in the gene encoding tyrosinase which is a key enzyme in melanin formation. However, exposure to UV radiation can increase melanin production, hence

931-467: Is strictly linked to seasonality. Traditionally, land preparation was done by hand. Nowadays, tractor plowing is also used. As maca grows on sites where no other crops can be cultivated, it is often found after long fallows of sheep grazing pastures. Maca croplands, thus, traditionally are only fertilized with sheep and alpaca manure; however, fertilizer application could prevent soils from depleting in nutrients . Weeding or pesticide application usually

980-412: Is the hypothesized pre-cellular stage in the evolutionary history of life on earth, in which self-replicating RNA molecules proliferated prior to the evolution of DNA and proteins. The folded three-dimensional physical structure of the first RNA molecule that possessed ribozyme activity promoting replication while avoiding destruction would have been the first phenotype, and the nucleotide sequence of

1029-701: The 1990s to 2014. Market studies showed low acceptance of the particular maca taste by consumers when first exposed to it, creating a barrier for popularity of this food as a culinary vegetable. The economic interest existed more in the perceived but unproven health effects of the root's constituents supplied as an extract in a dietary supplement. By 2014, agricultural and market interest in maca grew in China, but with challenges from Peruvian institutions who accused Chinese companies of illegally exporting maca and of biopiracy , as several Chinese patents had been filed to improve maca's propagation and genetic diversity. Maca

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1078-483: The color differences. Maca hypocotyls may be gold or cream, red, purple, blue, black, or green. Each is considered a "genetically unique variety," as the seeds of the parent plants grow to have roots of the same color. Specific phenotypes (in maca, 'phenotype' pertains mainly to root color) have been propagated exclusively to increase commercial interest. Cream-colored roots are the most widely grown and are favored in Peru for their enhanced sweetness and size. Black maca

1127-450: The concept of the phenotype has hidden subtleties. It may seem that anything dependent on the genotype is a phenotype, including molecules such as RNA and proteins . Most molecules and structures coded by the genetic material are not visible in the appearance of an organism, yet they are observable (for example by Western blotting ) and are thus part of the phenotype; human blood groups are an example. It may seem that this goes beyond

1176-517: The context of phenotype prediction. Although a phenotype is the ensemble of observable characteristics displayed by an organism, the word phenome is sometimes used to refer to a collection of traits, while the simultaneous study of such a collection is referred to as phenomics . Phenomics is an important field of study because it can be used to figure out which genomic variants affect phenotypes which then can be used to explain things like health, disease, and evolutionary fitness. Phenomics forms

1225-483: The corresponding amino acid sequence of a gene may change the frequency of guanine - cytosine base pairs ( GC content ). These base pairs have a higher thermal stability ( melting point ) than adenine - thymine , a property that might convey, among organisms living in high-temperature environments, a selective advantage on variants enriched in GC content. Richard Dawkins described a phenotype that included all effects that

1274-677: The cultivated maca L. meyenii . The Latin name recognized by the USDA similarly continues to be Lepidium meyenii . It has been debated whether it is botanically correct to consider meyenii and peruvianum to be distinct from one another. A 2015 multi-center study found differences in taxonomy, visual appearance, phytochemical profiles and DNA sequences when comparing L. meyenii and L. peruvianum , suggesting that they are different and that their names should not be considered synonyms. The growth habit, size, and proportions of maca are roughly similar to those of radishes and turnips , to which it

1323-572: The cultivated plants, maca is one of the most frost tolerant. Strong winds and sunlight are also characteristics of the native habitat of the maca. Maca today is still mainly cultivated in Peru , in the high Andes of Bolivia, and to a small extent also in Brazil . Maca can be cultivated beyond its natural elevation range, over 4,400 m (14,400 ft) above sea level. Maca (Peruvian ginseng) seedlings usually emerge about one month after sowing, with

1372-415: The environment plays a role in this phenotype as well. For most complex phenotypes the precise genetic mechanism remains unknown. For instance, it is largely unclear how genes determine the shape of bones or the human ear. Gene expression plays a crucial role in determining the phenotypes of organisms. The level of gene expression can affect the phenotype of an organism. For example, if a gene that codes for

1421-709: The evidence for the effectiveness of maca as a treatment for menopausal symptoms in women inconclusive. Phenotypes Wilhelm Johannsen proposed the genotype–phenotype distinction in 1911 to make clear the difference between an organism's hereditary material and what that hereditary material produces. The distinction resembles that proposed by August Weismann (1834–1914), who distinguished between germ plasm (heredity) and somatic cells (the body). More recently, in The Selfish Gene (1976), Dawkins distinguished these concepts as replicators and vehicles. Despite its seemingly straightforward definition,

1470-439: The evolution from genotype to genome to pan-genome , a concept of exploring the relationship ultimately among pan-phenome, pan-genome , and pan- envirome was proposed in 2023. Phenotypic variation (due to underlying heritable genetic variation ) is a fundamental prerequisite for evolution by natural selection . It is the living organism as a whole that contributes (or not) to the next generation, so natural selection affects

1519-707: The extracts of maca indicate a potential safety issue as a monoamine oxidase inhibitor (re. which see β-carboline ) and possibility as a mutagen . Due to these potential mutagenic properties of MTCA, the Agency for Sanitary Security in France warned consumers about the possible health risks of powdered maca root, a declaration disputed on the assumption that MTCA would be deactivated by boiling to process maca roots. MTCA-like compounds are associated with craving behaviour. Due to its purported effect on fertility , maca grew in agricultural, commercial, and research interest from

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1568-440: The false statement that a "mutation has no phenotype". Behaviors and their consequences are also phenotypes, since behaviors are observable characteristics. Behavioral phenotypes include cognitive, personality, and behavioral patterns. Some behavioral phenotypes may characterize psychiatric disorders or syndromes. A phenome is the set of all traits expressed by a cell , tissue , organ , organism , or species . The term

1617-541: The genetic structure of a population indirectly via the contribution of phenotypes. Without phenotypic variation, there would be no evolution by natural selection. The interaction between genotype and phenotype has often been conceptualized by the following relationship: A more nuanced version of the relationship is: Genotypes often have much flexibility in the modification and expression of phenotypes; in many organisms these phenotypes are very different under varying environmental conditions. The plant Hieracium umbellatum

1666-721: The individual. Large-scale genetic screens can identify the genes or mutations that affect the phenotype of an organism. Analyzing the phenotypes of mutant genes can also aid in determining gene function. Most genetic screens have used microorganisms, in which genes can be easily deleted. For instance, nearly all genes have been deleted in E. coli and many other bacteria , but also in several eukaryotic model organisms such as baker's yeast and fission yeast . Among other discoveries, such studies have revealed lists of essential genes . More recently, large-scale phenotypic screens have also been used in animals, e.g. to study lesser understood phenotypes such as behavior . In one screen,

1715-532: The late 1980s. It is grown for its fleshy hypocotyl that is fused with a taproot , which is typically dried but may also be freshly cooked as a root vegetable . As a cash crop, it is primarily exported as a powder that may be raw or processed further as a gelatinized starch or as an extract . If dried, it may be processed into a flour for baking or as a dietary supplement . Its Spanish and Quechua names include maca-maca , maino , ayak chichira , and ayak willku . Antonio Vázquez de Espinosa described

1764-626: The levels of gene expression can be influenced by a variety of factors, such as environmental conditions, genetic variations, and epigenetic modifications. These modifications can be influenced by environmental factors such as diet, stress, and exposure to toxins, and can have a significant impact on an individual's phenotype. Some phenotypes may be the result of changes in gene expression due to these factors, rather than changes in genotype. An experiment involving machine learning methods utilizing gene expressions measured from RNA sequencing found that they can contain enough signal to separate individuals in

1813-452: The maca an annual plant, as in favorable years, it can complete a lifecycle within a year. Maca is the only member of the genus Lepidium with a fleshy hypocotyl , which is fused with the taproot to form a rough inverted pear-shaped body. Traditionally, native growers have acknowledged four varieties of maca based on their root color: cream-yellow, half-purple, purple, and black; varying levels of anthocyanin are primarily responsible for

1862-619: The number of putative mutants (see table for details). Putative mutants are then tested for heritability in order to help determine the inheritance pattern as well as map out the mutations. Once they have been mapped out, cloned, and identified, it can be determined whether a mutation represents a new gene or not. These experiments showed that mutations in the rhodopsin gene affected vision and can even cause retinal degeneration in mice. The same amino acid change causes human familial blindness , showing how phenotyping in animals can inform medical diagnostics and possibly therapy. The RNA world

1911-466: The onset of the rainy season in October. In the vegetative phase, from May to June, the lower part of the hypocotyl, as well as the upper part of the tap root, grows in size. After 260 to 280 days, it is formed into the harvestable hypocotyl. If the root is left in the soil, it is dormant for two to three months in the time of the cold, dry season until August. Then, it will form a generative shoot on which

1960-411: The original intentions of the concept with its focus on the (living) organism in itself. Either way, the term phenotype includes inherent traits or characteristics that are observable or traits that can be made visible by some technical procedure. The term "phenotype" has sometimes been incorrectly used as a shorthand for the phenotypic difference between a mutant and its wild type , which would lead to

2009-438: The phenome of a given organism is best understood as a kind of matrix of data representing physical manifestation of phenotype. For example, discussions led by A. Varki among those who had used the term up to 2003 suggested the following definition: "The body of information describing an organism's phenotypes, under the influences of genetic and environmental factors". Another team of researchers characterize "the human phenome [as]

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2058-531: The phenotype that grows. An example of random variation in Drosophila flies is the number of ommatidia , which may vary (randomly) between left and right eyes in a single individual as much as they do between different genotypes overall, or between clones raised in different environments. The concept of phenotype can be extended to variations below the level of the gene that affect an organism's fitness. For example, silent mutations that do not change

2107-439: The plant following his visit to Peru circa 1598, and Bernabé Cobo described this plant in the early 17th century. Gerhard Walpers named the species Lepidium meyenii in 1843. In the 1990s, Gloria Chacon further distinguished a different species. She considered the widely cultivated natural maca of today to be a newer domesticated species, L. peruvianum . However, most botanists doubt this distinction and continue to call

2156-428: The role of mutations in mice were studied in areas such as learning and memory , circadian rhythmicity , vision, responses to stress and response to psychostimulants . This experiment involved the progeny of mice treated with ENU , or N-ethyl-N-nitrosourea, which is a potent mutagen that causes point mutations . The mice were phenotypically screened for alterations in the different behavioral domains in order to find

2205-442: The seeds ripen five months later. One plant can form up to 1000 tiny seeds, 1600 of which weigh about one gram. Thus, only relatively few plants are needed for propagation. The cultivated plants are selected for preferred size and color, then placed 50–100 mm deep in pits with alternate layers of grass and soil to protect them from drying out. They are fertilized heavily, as maca is an exhaustive crop of soil. The cultivation cycle

2254-540: The tough fiber from the roots using gentle heat and pressure, as raw maca is difficult to digest due to its thick fibers. Archaeological evidence exists for varying degrees of cultivation of maca in the Lake Junin region from around 1700 BC to 1200 AD. Maca has been harvested and used by humans in the puna grasslands of the Andean Mountains for centuries. Contrary to frequent claims that maca cultivation

2303-406: The vicinity of the growers. The root can also be mashed and boiled to produce a sweet, thick liquid, then dried and mixed with milk to form a porridge, mazamorra . The cooked roots are also used with other vegetables in empanadas , jams, or soups. The root may be ground to produce flour for bread, cakes, or pancakes. If fermented, a weak beer called chicha de maca may be produced. In 2010,

2352-594: Was common in what is today Peru, until the late 1980s, maca has been cultivated only in a limited area around Lake Junin in central Peru. Historically, maca was often traded for lowland tropical food staples, such as maize , rice , manioc (tapioca roots), quinoa , and papaya . It also was used as a form of payment for Spanish imperial taxes. Maca is said to have medicinal properties, but scientific research remains inconclusive. A 2016 systematic review found evidence suggesting that it improves semen quality in healthy and infertile men. A 2011 systematic review found

2401-405: Was first used by Davis in 1949, "We here propose the name phenome for the sum total of extragenic, non-autoreproductive portions of the cell, whether cytoplasmic or nuclear. The phenome would be the material basis of the phenotype, just as the genome is the material basis of the genotype ." Although phenome has been in use for many years, the distinction between the use of phenome and phenotype

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