Myco-heterotrophy (from Greek μύκης mykes, "fungus", ἕτερος heteros, "another", "different" and τροφή trophe, "nutrition") is a symbiotic relationship between certain kinds of plants and fungi, in which the plant gets all or part of its food from parasitism upon fungi rather than from photosynthesis. A myco-heterotroph is the parasitic plant partner in this relationship. Myco-heterotrophy is considered a kind of cheating relationship and myco-heterotrophs are sometimes informally referred to as "mycorrhizal cheaters". This relationship is sometimes referred to as mycotrophy, though this term is also used for plants that engage in mutualistic mycorrhizal relationships.
Full (or obligate) myco-heterotrophy exists when a non-photosynthetic plant (a plant largely lacking in chlorophyll or otherwise lacking a functional photosystem) gets all of its food from the fungi that it parasitizes. Partial (or facultative) myco-heterotrophy exists when a plant is capable of photosynthesis, but parasitizes fungi as a supplementary food supply. There are also plants, such as some orchid species, that are non-photosynthetic and obligately myco-heterotrophic for part of their life cycle, and photosynthetic and facultatively myco-heterotrophic or non-myco-heterotrophic for the rest of their life cycle. Not all non-photosynthetic or "achlorophyllous" plants are myco-heterotrophic – some non-photosynthetic plants like dodder directly parasitize the vascular tissue of other plants.
In the past, non-photosynthetic plants were mistakenly thought to get food by breaking down organic matter in a manner similar to saprotrophic fungi. Such plants were therefore called "saprophytes". It is now known that these plants are not physiologically capable of directly breaking down organic matter and that in order to get food, non-photosynthetic plants must engage in parasitism, either through myco-heterotrophy or direct parasitism of other plants.
The interface between the plant and fungal partners in this association is between the roots of the plant and the mycelium of the fungus. Myco-heterotrophy therefore closely resembles mycorrhiza (and indeed is thought to have evolved from mycorrhiza), except that in myco-heterotrophy, the flow of carbon is from the fungus to the plant, rather than vice versa.
Most myco-heterotrophs can therefore be seen as ultimately being epiparasites, since they take energy from fungi that in turn get their energy from vascular plants. Indeed, much myco-heterotrophy takes place in the context of common mycorrhizal networks, in which plants use mycorrhizal fungi to exchange carbon and nutrients with other plants. In these systems, myco-heterotrophs play the role of "mycorrhizal cheaters", taking carbon from the common network, with no known reward.
In congruence with older reports, it has been recently shown that some myco-heterotrophic orchids can be supported by saprotrophic fungi, exploiting litter- or wood-decaying fungi. In addition, several green plants (evolutionarily close to myco-heterotrophic species) have been shown to engage in partial myco-heterotrophy, that is, they are able to take carbon from mycorrhizal fungi, in addition to their photosynthetic intake.
Myco-heterotrophs are found among a number of plant groups. All monotropes and non-photosynthetic orchids are full myco-heterotrophs, as is the non-photosynthetic liverwort Cryptothallus. Partial myco-heterotrophy is common in the Gentian family, with a few genera such as Voyria being fully myco-heterotrophic; in photosynthetic orchids; and in a number of other plant groups. Some ferns and clubmosses have myco-heterotrophic gametophyte stages. The fungi that are parasitized by myco-heterotrophs are typically fungi with large energy reserves to draw on, usually mycorrhizal fungi, though there is some evidence that they may also parasitize parasitic fungi that form extensive mycelial networks, such as Armillaria. Examples of fungi parasitized by myco-heterotrophic plants can be found among the ectomycorrhizal, arbuscular mycorrhizal, and orchid mycorrhizal fungi. The great diversity in unrelated plant families with myco-heterotrophic members, as well as the diversity of fungi targeted by myco-heterotrophs, suggests multiple parallel evolution of myco-heterotrophs from mycorrhizal ancestors.
Burmanniaceae is a family of flowering plants, consisting of 99 species of herbaceous plants in eight genera.Cascade effect (ecology)
An ecological cascade effect is a series of secondary extinctions that is triggered by the primary extinction of a key species in an ecosystem. Secondary extinctions are likely to occur when the threatened species are: dependent on a few specific food sources, mutualistic (dependent on the key species in some way), or forced to coexist with an invasive species that is introduced to the ecosystem. Species introductions to a foreign ecosystem can often devastate entire communities, and even entire ecosystems. These exotic species monopolize the ecosystem's resources, and since they have no natural predators to decrease their growth, they are able to increase indefinitely. Olsen et al. showed that exotic species have caused lake and estuary ecosystems to go through cascade effects due to loss of algae, crayfish, mollusks, fish, amphibians, and birds. However, the principal cause of cascade effects is the loss of top predators as the key species. As a result of this loss, a dramatic increase (ecological release) of prey species occurs. The prey is then able to overexploit its own food resources, until the population numbers decrease in abundance, which can lead to extinction. When the prey's food resources disappear, they starve and may go extinct as well. If the prey species is herbivorous, then their initial release and exploitation of the plants may result in a loss of plant biodiversity in the area. If other organisms in the ecosystem also depend upon these plants as food resources, then these species may go extinct as well. An example of the cascade effect caused by the loss of a top predator is apparent in tropical forests. When hunters cause local extinctions of top predators, the predators' prey's population numbers increase, causing an overexploitation of a food resource and a cascade effect of species loss. Recent studies have been performed on approaches to mitigate extinction cascades in food-web networks.Corallorhiza
Corallorhiza, the coralroot, is a genus of flowering plants in the orchid family. Except for the circumboreal C. trifida, the genus is restricted to North America (including Mexico, Central America and the West Indies).Most species are putatively parasitic, relying entirely upon mycorrhizal fungi within their coral-shaped rhizomes for sustenance. Because of this dependence on myco-heterotrophy, they have never been successfully cultivated. Most species are leafless and rootless. Most species produce little or no chlorophyll, and do not utilize photosynthesis. An exception is the yellowish green species Corallorhiza trifida, which has some chlorophyll and is able to fix CO2. However, this species also depends primarily on fungal associations for carbon acquisition.Energy flow (ecology)
In ecology, energy flow, also called the calorific flow, refers to the flow of energy through a food chain, and is the focus of study in ecological energetics. In an ecosystem, ecologists seek to quantify the relative importance of different component species and feeding relationships.
A general energy flow scenario follows:
Solar energy is fixed by the photoautotrophs, called primary producers, like green plants. Primary consumers absorb most of the stored energy in the plant through digestion, and transform it into the form of energy they need, such as adenosine triphosphate (ATP), through respiration. A part of the energy received by primary consumers, herbivores, is converted to body heat (an effect of respiration), which is radiated away and lost from the system. The loss of energy through body heat is far greater in warm-blooded animals, which must eat much more frequently than those that are cold-blooded. Energy loss also occurs in the expulsion of undigested food (egesta) by excretion or regurgitation.
Secondary consumers, carnivores, then consume the primary consumers, although omnivores also consume primary producers. Energy that had been used by the primary consumers for growth and storage is thus absorbed into the secondary consumers through the process of digestion. As with primary consumers, secondary consumers convert this energy into a more suitable form (ATP) during respiration. Again, some energy is lost from the system, since energy which the primary consumers had used for respiration and regulation of body temperature cannot be utilized by the secondary consumers.
Tertiary consumers, which may or may not be apex predators, then consume the secondary consumers, with some energy passed on and some lost, as with the lower levels of the food chain.
A final link in the food chain are decomposers which break down the organic matter of the tertiary consumers (or whichever consumer is at the top of the chain) and release nutrients into the soil. They also break down plants, herbivores and carnivores that were not eaten by organisms higher on the food chain, as well as the undigested food that is excreted by herbivores and carnivores. Saprotrophic bacteria and fungi are decomposers, and play a pivotal role in the nitrogen and carbon cycles.The energy is passed on from trophic level to trophic level and each time about 90% of the energy is lost, with some being lost as heat into the environment (an effect of respiration) and some being lost as incompletely digested food (egesta). Therefore, primary consumers get about 10% of the energy produced by autotrophs, while secondary consumers get 1% and tertiary consumers get 0.1%. This means the top consumer of a food chain receives the least energy, as a lot of the food chain's energy has been lost between trophic levels. This loss of energy at each level limits typical food chains to only four to six links.Food web
A food web (or food cycle) is the natural interconnection of food chains and a graphical representation (usually an image) of what-eats-what in an ecological community. Another name for food web is consumer-resource system. Ecologists can broadly lump all life forms into one of two categories called trophic levels: 1) the autotrophs, and 2) the heterotrophs. To maintain their bodies, grow, develop, and to reproduce, autotrophs produce organic matter from inorganic substances, including both minerals and gases such as carbon dioxide. These chemical reactions require energy, which mainly comes from the Sun and largely by photosynthesis, although a very small amount comes from hydrothermal vents and hot springs. A gradient exists between trophic levels running from complete autotrophs that obtain their sole source of carbon from the atmosphere, to mixotrophs (such as carnivorous plants) that are autotrophic organisms that partially obtain organic matter from sources other than the atmosphere, and complete heterotrophs that must feed to obtain organic matter. The linkages in a food web illustrate the feeding pathways, such as where heterotrophs obtain organic matter by feeding on autotrophs and other heterotrophs. The food web is a simplified illustration of the various methods of feeding that links an ecosystem into a unified system of exchange. There are different kinds of feeding relations that can be roughly divided into herbivory, carnivory, scavenging and parasitism. Some of the organic matter eaten by heterotrophs, such as sugars, provides energy. Autotrophs and heterotrophs come in all sizes, from microscopic to many tonnes - from cyanobacteria to giant redwoods, and from viruses and bdellovibrio to blue whales.
Charles Elton pioneered the concept of food cycles, food chains, and food size in his classical 1927 book "Animal Ecology"; Elton's 'food cycle' was replaced by 'food web' in a subsequent ecological text. Elton organized species into functional groups, which was the basis for Raymond Lindeman's classic and landmark paper in 1942 on trophic dynamics. Lindeman emphasized the important role of decomposer organisms in a trophic system of classification. The notion of a food web has a historical foothold in the writings of Charles Darwin and his terminology, including an "entangled bank", "web of life", "web of complex relations", and in reference to the decomposition actions of earthworms he talked about "the continued movement of the particles of earth". Even earlier, in 1768 John Bruckner described nature as "one continued web of life".
Food webs are limited representations of real ecosystems as they necessarily aggregate many species into trophic species, which are functional groups of species that have the same predators and prey in a food web. Ecologists use these simplifications in quantitative (or mathematical representation) models of trophic or consumer-resource systems dynamics. Using these models they can measure and test for generalized patterns in the structure of real food web networks. Ecologists have identified non-random properties in the topographic structure of food webs. Published examples that are used in meta analysis are of variable quality with omissions. However, the number of empirical studies on community webs is on the rise and the mathematical treatment of food webs using network theory had identified patterns that are common to all. Scaling laws, for example, predict a relationship between the topology of food web predator-prey linkages and levels of species richness.Fungivore
Fungivory or mycophagy is the process of organisms consuming fungi. Many different organisms have been recorded to gain their energy from consuming fungi, including birds, mammals, insects, plants, amoebas, gastropods, nematodes, bacteria and other fungi. Some of these, which only eat fungi, are called fungivores whereas others eat fungi as only part of their diet, being omnivores.Fungus
A fungus (plural: fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, fungi, which is separate from the other eukaryotic life kingdoms of plants and animals.
A characteristic that places fungi in a different kingdom from plants, bacteria, and some protists is chitin in their cell walls. Similar to animals, fungi are heterotrophs; they acquire their food by absorbing dissolved molecules, typically by secreting digestive enzymes into their environment. Fungi do not photosynthesize. Growth is their means of mobility, except for spores (a few of which are flagellated), which may travel through the air or water. Fungi are the principal decomposers in ecological systems. These and other differences place fungi in a single group of related organisms, named the Eumycota (true fungi or Eumycetes), which share a common ancestor (form a monophyletic group), an interpretation that is also strongly supported by molecular phylogenetics. This fungal group is distinct from the structurally similar myxomycetes (slime molds) and oomycetes (water molds). The discipline of biology devoted to the study of fungi is known as mycology (from the Greek μύκης mykes, mushroom). In the past, mycology was regarded as a branch of botany, although it is now known fungi are genetically more closely related to animals than to plants.
Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their cryptic lifestyles in soil or on dead matter. Fungi include symbionts of plants, animals, or other fungi and also parasites. They may become noticeable when fruiting, either as mushrooms or as molds. Fungi perform an essential role in the decomposition of organic matter and have fundamental roles in nutrient cycling and exchange in the environment. They have long been used as a direct source of human food, in the form of mushrooms and truffles; as a leavening agent for bread; and in the fermentation of various food products, such as wine, beer, and soy sauce. Since the 1940s, fungi have been used for the production of antibiotics, and, more recently, various enzymes produced by fungi are used industrially and in detergents. Fungi are also used as biological pesticides to control weeds, plant diseases and insect pests. Many species produce bioactive compounds called mycotoxins, such as alkaloids and polyketides, that are toxic to animals including humans. The fruiting structures of a few species contain psychotropic compounds and are consumed recreationally or in traditional spiritual ceremonies. Fungi can break down manufactured materials and buildings, and become significant pathogens of humans and other animals. Losses of crops due to fungal diseases (e.g., rice blast disease) or food spoilage can have a large impact on human food supplies and local economies.
The fungus kingdom encompasses an enormous diversity of taxa with varied ecologies, life cycle strategies, and morphologies ranging from unicellular aquatic chytrids to large mushrooms. However, little is known of the true biodiversity of Kingdom Fungi, which has been estimated at 2.2 million to 3.8 million species. Of these, only about 120,000 have been described, with over 8,000 species known to be detrimental to plants and at least 300 that can be pathogenic to humans. Ever since the pioneering 18th and 19th century taxonomical works of Carl Linnaeus, Christian Hendrik Persoon, and Elias Magnus Fries, fungi have been classified according to their morphology (e.g., characteristics such as spore color or microscopic features) or physiology. Advances in molecular genetics have opened the way for DNA analysis to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits. Phylogenetic studies published in the last decade have helped reshape the classification within Kingdom Fungi, which is divided into one subkingdom, seven phyla, and ten subphyla.Gentianaceae
Gentianaceae is a family of flowering plants of 87 genera and about 1600 species.Invasive species
An invasive species is a species that is not native to a specific location (an introduced species), and that has a tendency to spread to a degree believed to cause damage to the environment, human economy or human health.The term as most often used applies to introduced species that adversely affect the habitats and bioregions they invade economically, environmentally, or ecologically. Such species may be either plants or animals and may disrupt by dominating a region, wilderness areas, particular habitats, or wildland–urban interface land from loss of natural controls (such as predators or herbivores). This includes plant species labeled as exotic pest plants and invasive exotics growing in native plant communities. The European Union defines "Invasive Alien Species" as those that are, firstly, outside their natural distribution area, and secondly, threaten biological diversity. The term is also used by land managers, botanists, researchers, horticulturalists, conservationists, and the public for noxious weeds.The term "invasive" is often poorly defined or very subjective and some broaden the term to include indigenous or "native" species, that have colonized natural areas - for example deer considered by some to be overpopulating their native zones and adjacent suburban gardens in the Northeastern and Pacific Coast regions of the United States.The definition of "native" is also sometimes controversial. For example, the ancestors of Equus ferus (modern horses) evolved in North America and radiated to Eurasia before becoming locally extinct. Upon returning to North America in 1493 during their hominid-assisted migration, it is debatable as to whether they were native or exotic to the continent of their evolutionary ancestors.Notable examples of invasive plant species include The kudzu vine, Andean pampas grass, and yellow starthistle. Animal examples include the New Zealand mud snail, feral pigs, European rabbits, grey squirrels, domestic cats, carp and ferrets.Invasion of long-established ecosystems by organisms from distant bio-regions is a natural phenomenon, but has been accelerated massively by humans, from their earliest migrations though to the age of discovery, and now international trade.Lecanorchis tabugawaensis
Lecanorchis tabugawaensis is a species of orchid which belongs to the group of mycoheterotrophic plants. Mycoheterotrophs are known for feeding off of the roots of fungal hosts, instead of utilizing photosynthesis. Myco-heterotrophy refers to the harmonious relationship between particular plants and fungi. After a thorough exploration of the plant's shape, size, and structure, it was determined to be closely associated with orchidaceous Lecanorchis amethystea, although inward variations confirmed this unique species.
The plant was discovered in July 2015 by Yamashita Hiroaki in lowland forests on Yakushima island. This species is only found at two locations along the Tabu River. Other recent discoveries on this island include: Oxygyne yamashitae (2008), Gastrodia uraiensis (2015), and Sciaphila yakushimensis (2016). This island also harbors various scarce plant species such as Lecanorchis virella, Lecanorchis trachycaula, Yakushimense vexillabium, Apostoasia nipponica, Lycopodium sieboldii, and Lysionotus pauciflorus.Mesopredator release hypothesis
The mesopredator release hypothesis is an ecological theory used to describe the interrelated population dynamics between apex predators and mesopredators within an ecosystem, such that a collapsing population of the former results in dramatically-increased populations of the latter. This hypothesis describes the phenomenon of trophic cascade in specific terrestrial communities.
A mesopredator is a medium-sized, middle trophic level predator, which both preys and is preyed upon. Examples are raccoons, skunks, snakes, cownose rays, and small sharks.Monotropoideae
Monotropoideae, sometimes referred to as monotropes, are a flowering plant subfamily in the family Ericaceae. Members of this subfamily are notable for their mycoheterotrophic and non-photosynthesizing or achlorophyllous characteristics.Mycotroph
A mycotroph is a plant that gets all or part of its carbon, water, or nutrient supply through symbiotic association with fungi. The term can refer to plants that engage in either of two distinct symbioses with fungi:
Many mycotrophs have a mutualistic association with fungi in any of several forms of mycorrhiza. The majority of plant species are mycotrophic in this sense. Examples include Burmanniaceae.
Some mycotrophs are parasitic upon fungi in an association known as myco-heterotrophy.Piperia yadonii
Piperia yadonii, also known as Yadon's piperia or Yadon's rein orchid, is an endangered orchid endemic to a narrow range of coastal habitat in northern Monterey County, California. In 1998 this plant was designated as an endangered species by the United States government, the major threat to its survival being continuing land development from an expanding human population and associated habitat loss. One of the habitats of Yadon's Piperia, the Del Monte Forest near Monterey, California, is the subject of a federal lawsuit, based upon endangerment of this organism along with several other endangered species.
This wildflower may lie dormant in a given year and not emerge above the soil surface from its tuberous substructure. After leafing out in the spring, it will produce flowers on erect spikes, each flower possessing both green and white petals. It prefers sandy soils, and subsists on nutrient extraction from intermediate fungal organisms.Productivity (ecology)
In ecology, productivity refers to the rate of generation of biomass in an ecosystem. It is usually expressed in units of mass per unit surface (or volume) per unit time, for instance grams per square metre per day (g m−2 d−1). The mass unit may relate to dry matter or to the mass of carbon generated. Productivity of autotrophs such as plants is called primary productivity, while that of heterotrophs such as animals is called secondary productivity.Russulaceae
The Russulaceae are a diverse family of fungi in the order Russulales, with roughly 1,900 known species and a worldwide distribution. They comprise the brittlegills and the milk-caps, well-known mushroom-forming fungi that include some edible species. These gilled mushrooms are characterised by the brittle flesh of their fruitbodies.
In addition to these typical agaricoid forms, the family contains species with fruitbodies that are laterally striped (pleurotoid), closed (secotioid or gasteroid), or crust-like (corticioid). Molecular phylogenetics has demonstrated close affinities between species with very different fruitbody types and has discovered new, distinct lineages.
An important group of root-symbiotic ectomycorrhizal fungi in forests and shrublands around the world includes Lactifluus, Multifurca, Russula, and Lactarius. The crust-forming genera Boidinia, Gloeopeniophorella, and Pseudoxenasma, all wood-decay fungi, have basal positions in the family.Sustainable gardening
Sustainable gardening includes the more specific sustainable landscapes, sustainable landscape design, sustainable landscaping, sustainable landscape architecture, resulting in sustainable sites. It comprises a disparate group of horticultural interests that can share the aims and objectives associated with the international post-1980s sustainable development and sustainability programs developed to address the fact that humans are now using natural biophysical resources faster than they can be replenished by nature.Included within this compass are those home gardeners, and members of the landscape and nursery industries, and municipal authorities, that integrate environmental, social, and economic factors to create a more sustainable future.
Organic gardening and the use of native plants are integral to sustainable gardening.The Phytologist
The Phytologist was a British botanical journal, appearing first as Phytologist: a popular botanical miscellany. It was founded in 1841 as a monthly, edited by George Luxford. Luxford died in 1854, and the title was taken over by Alexander Irvine and William Pamplin, who ran it to 1863 with subtitle "a botanical journal".The proprietor for the first series was Edward Newman, also a contributor. The publisher was John Van Voorst. The journal never made money. Newman used its pages to attack Vestiges of Creation (1844), in an outspoken signed review that stood out from the mass of anonymous comment. Luxford's overall editorial policy, however, gave space to those supporting transmutation of species. The Phytologist, quite unofficially, became the house journal of the Botanical Society of London; and Hewett Watson of the Society a prominent contributor. In the early issues Luxford wrote a series of ten articles on myco-heterotrophy, around Monotropa hypopithys, and prompted sharp debate.