Symbiosis

From Wikipedia, the free encyclopedia

In a cleaning symbiosis, the clownfish feeds on small invertebrates that otherwise have potential to harm the sea anemone, and the fecal matter from the clownfish provides nutrients to the sea anemone. The clownfish is protected from predators by the anemone's stinging cells, to which the clownfish is immune. The clownfish emits a high pitched sound that deters butterfly fish, which would otherwise eat the anemone, making the relationship appear mutualistic.

 

Symbiosis (from Greek συμβίωσις "living together", from σύν "together" and βίωσις "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasitic. The organisms, each termed a symbiont, may be of the same or of different species. In 1879, Heinrich Anton de Bary defined it as "the living together of unlike organisms". The term was subject to a century-long debate about whether it should specifically denote mutualism, as in lichens; biologists have now abandoned that restriction.

Symbiosis can be obligatory, which means that one or both of the symbionts entirely depend on each other for survival, or facultative (optional) when they can generally live independently. 

Symbiosis is also classified by physical attachment; symbiosis in which the organisms have bodily union is called conjunctive symbiosis, and symbiosis in which they are not in union is called disjunctive symbiosis. When one organism lives on the surface of another, such as head lice on humans, it is called ectosymbiosis; when one partner lives inside the tissues of another, such as Symbiodinium within coral, it is termed endosymbiosis.

Definition

Diagram of the six possible types of symbiotic relationship, from mutual benefit to mutual harm.

 

The definition of symbiosis was a matter of debate for 130 years. In 1877, Albert Bernhard Frank used the term symbiosis to describe the mutualistic relationship in lichens. In 1879, the German mycologist Heinrich Anton de Bary defined it as "the living together of unlike organisms". The definition has varied among scientists, with some advocating that it should only refer to persistent mutualisms, while others thought it should apply to all persistent biological interactions, in other words mutualisms, commensalism, or parasitism, but excluding brief interactions such as predation. Current biology and ecology textbooks use the latter "de Bary" definition, or an even broader one where symbiosis means all interspecific interactions; the restrictive definition where symbiosis means only mutualism is no longer used.

In 1949, Edward Haskell proposed an integrative approach, proposing a classification of "co-actions", later adopted by biologists as "interactions".

Biological interactions can involve individuals of the same species (intraspecific interactions) or individuals of different species (interspecific interactions). These can be further classified by either the mechanism of the interaction or the strength, duration and direction of their effects.

Obligate versus facultative

Relationships can be obligate, meaning that one or both of the symbionts entirely depend on each other for survival. For example, in lichens, which consist of fungal and photosynthetic symbionts, the fungal partners cannot live on their own. The algal or cyanobacterial symbionts in lichens, such as Trentepohlia, can generally live independently, and their symbiosis is, therefore, facultative (optional).

Physical interaction

Alder tree root nodule houses endosymbiotic nitrogen-fixing bacteria.

Endosymbiosis is any symbiotic relationship in which one symbiont lives within the tissues of the other, either within the cells or extracellularly. Examples include diverse microbiomes, rhizobia, nitrogen-fixing bacteria that live in root nodules on legume roots; actinomycete, nitrogen-fixing bacteria such as Frankia, which live in alder root nodules; single-celled algae inside reef-building corals; and bacterial endosymbionts that provide essential nutrients to about 10%–15% of insects.

Ectosymbiosis is any symbiotic relationship in which the symbiont lives on the body surface of the host, including the inner surface of the digestive tract or the ducts of exocrine glands. Examples of this include ectoparasites such as lice; commensal ectosymbionts such as the barnacles, which attach themselves to the jaw of baleen whales; and mutualist ectosymbionts such as cleaner fish. 

Male-male interference competition in red deer

Competition

Competition can be defined as an interaction between organisms or species, in which the fitness of one is lowered by the presence of another. Limited supply of at least one resource (such as food, water, and territory) used by both usually facilitates this type of interaction, although the competition may also exist over other 'amenities', such as females for reproduction (in case of male organisms of the same species).

Mutualism

Hermit crab, Calcinus laevimanus, with sea anemone.

 

Mutualism or interspecies reciprocal altruism is a long-term relationship between individuals of different species where both individuals benefit. Mutualistic relationships may be either obligate for both species, obligate for one but facultative for the other, or facultative for both. 

Bryoliths document a mutualistic symbiosis between a hermit crab and encrusting bryozoans.

 

A large percentage of herbivores have mutualistic gut flora to help them digest plant matter, which is more difficult to digest than animal prey. This gut flora is made up of cellulose-digesting protozoans or bacteria living in the herbivores' intestines. Coral reefs are the result of mutualisms between coral organisms and various types of algae which live inside them. Most land plants and land ecosystems rely on mutualisms between the plants, which fix carbon from the air, and mycorrhyzal fungi, which help in extracting water and minerals from the ground.

An example of mutualism is the relationship between the ocellaris clownfish that dwell among the tentacles of Ritteri sea anemones. The territorial fish protects the anemone from anemone-eating fish, and in turn the stinging tentacles of the anemone protect the clownfish from its predators. A special mucus on the clownfish protects it from the stinging tentacles.

A further example is the goby, a fish which sometimes lives together with a shrimp. The shrimp digs and cleans up a burrow in the sand in which both the shrimp and the goby fish live. The shrimp is almost blind, leaving it vulnerable to predators when outside its burrow. In case of danger, the goby touches the shrimp with its tail to warn it. When that happens both the shrimp and goby quickly retreat into the burrow. Different species of gobies (Elacatinus spp.) also clean up ectoparasites in other fish, possibly another kind of mutualism.

A non-obligate symbiosis is seen in encrusting bryozoans and hermit crabs. The bryozoan colony (Acanthodesia commensale) develops a cirumrotatory growth and offers the crab (Pseudopagurus granulimanus) a helicospiral-tubular extension of its living chamber that initially was situated within a gastropod shell.

Many types of tropical and sub-tropical ants have evolved very complex relationships with certain tree species.

Endosymbiosis

In endosymbiosis, the host cell lacks some of the nutrients which the endosymbiont provides. As a result, the host favors endosymbiont's growth processes within itself by producing some specialized cells. These cells affect the genetic composition of the host in order to regulate the increasing population of the endosymbionts and ensure that these genetic changes are passed onto the offspring via vertical transmission (heredity).

A spectacular example of obligate mutualism is the relationship between the siboglinid tube worms and symbiotic bacteria that live at hydrothermal vents and cold seeps. The worm has no digestive tract and is wholly reliant on its internal symbionts for nutrition. The bacteria oxidize either hydrogen sulfide or methane, which the host supplies to them. These worms were discovered in the late 1980s at the hydrothermal vents near the Galapagos Islands and have since been found at deep-sea hydrothermal vents and cold seeps in all of the world's oceans.

As the endosymbiont adapts to the host's lifestyle, the endosymbiont changes dramatically. There is a drastic reduction in its genome size, as many genes are lost during the process of metabolism, and DNA repair and recombination, while important genes participating in the DNA-to-RNA transcription, protein translation and DNA/RNA replication are retained. The decrease in genome size is due to loss of protein coding genes and not due to lessening of inter-genic regions or open reading frame (ORF) size. Species that are naturally evolving and contain reduced sizes of genes can be accounted for an increased number of noticeable differences between them, thereby leading to changes in their evolutionary rates. When endosymbiotic bacteria related with insects are passed on to the offspring strictly via vertical genetic transmission, intracellular bacteria go across many hurdles during the process, resulting in the decrease in effective population sizes, as compared to the free-living bacteria. The incapability of the endosymbiotic bacteria to reinstate their wild type phenotype via a recombination process is called Muller's ratchet phenomenon. Muller's ratchet phenomenon, together with less effective population sizes, leads to an accretion of deleterious mutations in the non-essential genes of the intracellular bacteria. This can be due to lack of selection mechanisms prevailing in the relatively "rich" host environment.

Commensalism

Commensal mites travelling (phoresy) on a fly (Pseudolynchia canariensis)

Commensalism describes a relationship between two living organisms where one benefits and the other is not significantly harmed or helped. It is derived from the English word commensal, used of human social interaction. It derives from a medieval Latin word meaning sharing food, formed from com- (with) and mensa (table).

Commensal relationships may involve one organism using another for transportation (phoresy) or for housing (inquilinism), or it may also involve one organism using something another created, after its death (metabiosis). Examples of metabiosis are hermit crabs using gastropod shells to protect their bodies, and spiders building their webs on plants.

Parasitism

Head (scolex) of tapeworm Taenia solium is adapted to parasitism with hooks and suckers to attach to its host.

 

In a parasitic relationship, the parasite benefits while the host is harmed. Parasitism takes many forms, from endoparasites that live within the host's body to ectoparasites and parasitic castrators that live on its surface and micropredators like mosquitoes that visit intermittently. Parasitism is an extremely successful mode of life; as many as half of all animals have at least one parasitic phase in their life cycles, and it is also frequent in plants and fungi. Moreover, almost all free-living animal species are hosts to parasites, often of more than one species.

Mimicry

Mimicry is a form of symbiosis in which a species adopts distinct characteristics of another species to alter its relationship dynamic with the species being mimicked, to its own advantage. Among the many types of mimicry are Batesian and Müllerian, the first involving one-sided exploitation, the second providing mutual benefit. Batesian mimicry is an exploitative three-party interaction where one species, the mimic, has evolved to mimic another, the model, to deceive a third, the dupe. In terms of signalling theory, the mimic and model have evolved to send a signal; the dupe has evolved to receive it from the model. This is to the advantage of the mimic but to the detriment of both the model, whose protective signals are effectively weakened, and of the dupe, which is deprived of an edible prey. For example, a wasp is a strongly-defended model, which signals with its conspicuous black and yellow coloration that it is an unprofitable prey to predators such as birds which hunt by sight; many hoverflies are Batesian mimics of wasps, and any bird that avoids these hoverflies is a dupe. In contrast, Müllerian mimicry is mutually beneficial as all participants are both models and mimics. For example, different species of bumblebee mimic each other, with similar warning coloration in combinations of black, white, red, and yellow, and all of them benefit from the relationship. 

Amensalism

The black walnut secretes a chemical from its roots that harms neighboring plants, an example of antagonism.

 

Amensalism is an asymmetric interaction where one species is harmed or killed by the other, and one is unaffected by the other. There are two types of amensalism, competition and antagonism (or antibiosis). Competition is where a larger or stronger organism deprives a smaller or weaker one from a resource. Antagonism occurs when one organism is damaged or killed by another through a chemical secretion. An example of competition is a sapling growing under the shadow of a mature tree. The mature tree can rob the sapling of necessary sunlight and, if the mature tree is very large, it can take up rainwater and deplete soil nutrients. Throughout the process, the mature tree is unaffected by the sapling. Indeed, if the sapling dies, the mature tree gains nutrients from the decaying sapling. An example of antagonism is Juglans nigra (black walnut), secreting juglone, a substance which destroys many herbaceous plants within its root zone.

A clear case of amensalism is where sheep or cattle trample grass. Whilst the presence of the grass causes negligible detrimental effects to the animal's hoof, the grass suffers from being crushed. Amensalism is often used to describe strongly asymmetrical competitive interactions, such as has been observed between the Spanish ibex and weevils of the genus Timarcha which feed upon the same type of shrub. Whilst the presence of the weevil has almost no influence on food availability, the presence of ibex has an enormous detrimental effect on weevil numbers, as they consume significant quantities of plant matter and incidentally ingest the weevils upon it.

Cleaning symbiosis

Cleaning symbiosis is an association between individuals of two species, where one (the cleaner) removes and eats parasites and other materials from the surface of the other (the client). It is putatively mutually beneficial, but biologists have long debated whether it is mutual selfishness, or simply exploitative. Cleaning symbiosis is well-known among marine fish, where some small species of cleaner fish, notably wrasses but also species in other genera, are specialised to feed almost exclusively by cleaning larger fish and other marine animals.

Co-evolution

Leafhoppers protected by meat ants

 

Symbiosis is increasingly recognized as an important selective force behind evolution; many species have a long history of interdependent co-evolution.

Symbiogenesis

Eukaryotes (plants, animals, fungi, and protists) developed by symbiogenesis from a symbiosis between bacteria and archaea. Evidence for this includes the fact that mitochondria and chloroplasts divide independently of the cell, and the observation that some organelles seem to have their own genome.

The biologist Lynn Margulis, famous for her work on endosymbiosis, contended that symbiosis is a major driving force behind evolution. She considered Darwin's notion of evolution, driven by competition, to be incomplete and claimed that evolution is strongly based on co-operation, interaction, and mutual dependence among organisms. According to Margulis and her son Dorion Sagan, "Life did not take over the globe by combat, but by networking."

Co-evolutionary relationships

Mycorrhizas

About 80% of vascular plants worldwide form symbiotic relationships with fungi, in particular in arbuscular mycorrhizas.

Pollination is a mutualism between flowering plants and their animal pollinators.

Pollination

A fig is pollinated by the fig wasp, Blastophaga psenes.

Flowering plants and the animals that pollinate them have co-evolved. Many plants that are pollinated by insects (in entomophily), bats, or birds (in ornithophily) have highly specialized flowers modified to promote pollination by a specific pollinator that is correspondingly adapted. The first flowering plants in the fossil record had relatively simple flowers. Adaptive speciation quickly gave rise to many diverse groups of plants, and, at the same time, corresponding speciation occurred in certain insect groups. Some groups of plants developed nectar and large sticky pollen, while insects evolved more specialized morphologies to access and collect these rich food sources. In some taxa of plants and insects, the relationship has become dependent, where the plant species can only be pollinated by one species of insect.

Pseudomyrmex ant on bull thorn acacia (Vachellia cornigera) with Beltian bodies that provide the ants with protein

Acacia ants and acacias

The acacia ant (Pseudomyrmex ferruginea) is an obligate plant ant that protects at least five species of "Acacia" (Vachellia) from preying insects and from other plants competing for sunlight, and the tree provides nourishment and shelter for the ant and its larvae.

Mycorrhiza

From Wikipedia, the free encyclopedia

Many conspicuous fungi such as the fly agaric (upper left) form ectomycorrhiza (upper right) with tree rootlets. Arbuscular mycorrhiza (lower left) are very common in plants, including crop species such as wheat (lower right)

 

A mycorrhiza (from Greek μύκης mýkēs, "fungus", and ῥίζα rhiza, "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant's rhizosphere, its root system. Mycorrhizae play important roles in plant nutrition, soil biology and soil chemistry.

In a mycorrhizal association, the fungus colonizes the host plant's root tissues, either intracellularly as in arbuscular mycorrhizal fungi (AMF or AM), or extracellularly as in ectomycorrhizal fungi. The association is sometimes mutualistic. In particular species or in particular circumstances mycorrhizae may have a parasitic association with host plants.

Definition

A mycorrhiza is a symbiotic association between a green plant and a fungus. The plant makes organic molecules such as sugars by photosynthesis and supplies them to the fungus, and the fungus supplies to the plant water and mineral nutrients, such as phosphorus, taken from the soil. Mycorrhizas are located in the roots of vascular plants, but mycorrhiza-like associations also occur in bryophytes and there is fossil evidence that early land plants that lacked roots formed arbuscular mycorrhizal associations. Most plant species form mycorrhizal associations, though some families like Brassicaceae and Chenopodiaceae cannot. Different forms for the association are detailed in the next section. The most common is the arbuscular type that is present in 70% of plant species, including many crop plants such as wheat and rice.

Types

Mycorrhizas are commonly divided into ectomycorrhizas and endomycorrhizas. The two types are differentiated by the fact that the hyphae of ectomycorrhizal fungi do not penetrate individual cells within the root, while the hyphae of endomycorrhizal fungi penetrate the cell wall and invaginate the cell membrane. Endomycorrhiza includes arbuscular, ericoid, and orchid mycorrhiza, while arbutoid mycorrhizas can be classified as ectoendomycorrhizas. Monotropoid mycorrhizas form a special category.

Ectomycorrhiza

Beech is ectomycorrhizal

 

Leccinum aurantiacum, an ectomycorrhizal fungus

Ectomycorrhizas, or EcM, are symbiotic associations between the roots of around 10% of plant families, mostly woody plants including the birch, dipterocarp, eucalyptus, oak, pine, and rose families, orchids, and fungi belonging to the Basidiomycota, Ascomycota, and Zygomycota. Some EcM fungi, such as many Leccinum and Suillus, are symbiotic with only one particular genus of plant, while other fungi, such as the Amanita, are generalists that form mycorrhizas with many different plants. An individual tree may have 15 or more different fungal EcM partners at one time. Thousands of ectomycorrhizal fungal species exist, hosted in over 200 genera. A recent study has conservatively estimated global ectomycorrhizal fungal species richness at approximately 7750 species, although, on the basis of estimates of knowns and unknowns in macromycete diversity, a final estimate of ECM species richness would probably be between 20,000 and 25,000.

Ectomycorrhizas consist of a hyphal sheath, or mantle, covering the root tip and a Hartig net of hyphae surrounding the plant cells within the root cortex. In some cases the hyphae may also penetrate the plant cells, in which case the mycorrhiza is called an ectendomycorrhiza. Outside the root, ectomycorrhizal extramatrical mycelium forms an extensive network within the soil and leaf litter. 

Nutrients can be shown to move between different plants through the fungal network. Carbon has been shown to move from paper birch trees into Douglas-fir trees thereby promoting succession in ecosystems. The ectomycorrhizal fungus Laccaria bicolor has been found to lure and kill springtails to obtain nitrogen, some of which may then be transferred to the mycorrhizal host plant. In a study by Klironomos and Hart, Eastern White Pine inoculated with L. bicolor was able to derive up to 25% of its nitrogen from springtails. When compared to non-mycorrhizal fine roots, ectomycorrhizae may contain very high concentrations of trace elements, including toxic metals (cadmium, silver) or chlorine.

The first genomic sequence for a representative of symbiotic fungi, the ectomycorrhizal basidiomycete L. bicolor, has been published. An expansion of several multigene families occurred in this fungus, suggesting that adaptation to symbiosis proceeded by gene duplication. Within lineage-specific genes those coding for symbiosis-regulated secreted proteins showed an up-regulated expression in ectomycorrhizal root tips suggesting a role in the partner communication. L. bicolor is lacking enzymes involved in the degradation of plant cell wall components (cellulose, hemicellulose, pectins and pectates), preventing the symbiont from degrading host cells during the root colonisation. By contrast, L. bicolor possesses expanded multigene families associated with hydrolysis of bacterial and microfauna polysaccharides and proteins. This genome analysis revealed the dual saprotrophic and biotrophic lifestyle of the mycorrhizal fungus that enables it to grow within both soil and living plant roots.

Arbutoid mycorrhiza

This type of mycorrhiza involves plants of the Ericaceae subfamily Arbutoideae. It is however different from ericoid mycorrhiza and resembles ectomycorrhiza, both functionally and in terms of the fungi involved. The difference to ectomycorrhiza is that some hyphae actually penetrate into the root cells, making this type of mycorrhiza an ectendomycorrhiza.

Wheat is arbuscular mycorrhizal

Endomycorrhiza

Arbuscular mycorrhiza

Endomycorrhizas are variable and have been further classified as arbuscular, ericoid, arbutoid, monotropoid, and orchid mycorrhizas. Arbuscular mycorrhizas, or AM (formerly known as vesicular-arbuscular mycorrhizas, or VAM), are mycorrhizas whose hyphae penetrate plant cells, producing structures that are either balloon-like (vesicles) or dichotomously branching invaginations (arbuscules) as a means of nutrient exchange. The fungal hyphae do not in fact penetrate the protoplast (i.e. the interior of the cell), but invaginate the cell membrane. The structure of the arbuscules greatly increases the contact surface area between the hypha and the cell cytoplasm to facilitate the transfer of nutrients between them. 

Arbuscular mycorrhizas are formed only by fungi in the division Glomeromycota. Fossil evidence and DNA sequence analysis suggest that this mutualism appeared 400-460 million years ago, when the first plants were colonizing land. Arbuscular mycorrhizas are found in 85% of all plant families, and occur in many crop species. The hyphae of arbuscular mycorrhizal fungi produce the glycoprotein glomalin, which may be one of the major stores of carbon in the soil. Arbuscular mycorrhizal fungi have (possibly) been asexual for many millions of years and, unusually, individuals can contain many genetically different nuclei (a phenomenon called heterokaryosis).

Ericoid mycorrhiza

An ericoid mycorrhizal fungus isolated from Woollsia pungens

Ericoid mycorrhizas are the third of the three more ecologically important types. They have a simple intraradical (grow in cells) phase, consisting of dense coils of hyphae in the outermost layer of root cells. There is no periradical phase and the extraradical phase consists of sparse hyphae that don't extend very far into the surrounding soil. They might form sporocarps (probably in the form of small cups), but their reproductive biology is little understood.

Ericoid mycorrhizas have also been shown to have considerable saprotrophic capabilities, which would enable plants to receive nutrients from not-yet-decomposed materials via the decomposing actions of their ericoid partners.

Orchid mycorrhiza

All orchids are myco-heterotrophic at some stage during their lifecycle and form orchid mycorrhizas with a range of basidiomycete fungi. Their hyphae penetrate into the root cells and form pelotons (coils) for nutrient exchange.

Monotropoid mycorrhiza

This type of mycorrhiza occurs in the subfamily Monotropoideae of the Ericaceae, as well as several genera in the Orchidaceae. These plants are heterotrophic or mixotrophic and derive their carbon from the fungus partner. This is thus a non-mutualistic, parasitic type of mycorrhizal symbiosis.

Mutualist dynamics

Mycorrhizal fungi form a mutualistic relationship with the roots of most plant species. In such a relationship, both the plants themselves and those parts of the roots that host the fungi, are said to be mycorrhizal. Relatively few of the mycorrhizal relationships between plant species and fungi have been examined to date, but 95% of the plant families investigated are predominantly mycorrhizal either in the sense that most of their species associate beneficially with mycorrhizae, or are absolutely dependent on mycorrhizae. The Orchidaceae are notorious as a family in which the absence of the correct mycorrhizae is fatal even to germinating seeds.

Recent research into ectomycorrhizal plants in boreal forests has indicated that mycorrhizal fungi and plants have a relationship that may be more complex than simply mutualistic. This relationship was noted when mycorrhizal fungi were unexpectedly found to be hoarding nitrogen from plant roots in times of nitrogen scarcity. Researchers argue that some mycorrhizae distribute nutrients based upon the environment with surrounding plants and other mycorrhizae. They go on to explain how this updated model could explain why mycorrhizae do not alleviate plant nitrogen limitation, and why plants can switch abruptly from a mixed strategy with both mycorrhizal and nonmycorrhizal roots to a purely mycorrhizal strategy as soil nitrogen availability declines. It has also been suggested that evolutionary and phylogenetic relationships can explain much more variation in the strength of mycorrhizal mutualisms than ecological factors.

Sugar-water/mineral exchange

In this mutualism, fungal hyphae (E) increase the surface area of the root and uptake of key nutrients while the plant supplies the fungi with fixed carbon (A=root cortex, B=root epidermis, C=arbuscle, D=vesicle, F=root hair, G=nuclei).

 

The mycorrhizal mutualistic association provides the fungus with relatively constant and direct access to carbohydrates, such as glucose and sucrose. The carbohydrates are translocated from their source (usually leaves) to root tissue and on to the plant's fungal partners. In return, the plant gains the benefits of the mycelium's higher absorptive capacity for water and mineral nutrients, partly because of the large surface area of fungal hyphae, which are much longer and finer than plant root hairs, and partly because some such fungi can mobilize soil minerals unavailable to the plants' roots. The effect is thus to improve the plant's mineral absorption capabilities.

Unaided plant roots may be unable to take up nutrients that are chemically or physically immobilised; examples include phosphate ions and micronutrients such as iron. One form of such immobilization occurs in soil with high clay content, or soils with a strongly basic pH. The mycelium of the mycorrhizal fungus can, however, access many such nutrient sources, and make them available to the plants they colonize. Thus, many plants are able to obtain phosphate, without using soil as a source. Another form of immobilisation is when nutrients are locked up in organic matter that is slow to decay, such as wood, and some mycorrhizal fungi act directly as decay organisms, mobilising the nutrients and passing some onto the host plants; for example, in some dystrophic forests, large amounts of phosphate and other nutrients are taken up by mycorrhizal hyphae acting directly on leaf litter, bypassing the need for soil uptake. Inga alley cropping, proposed as an alternative to slash and burn rainforest destruction, relies upon mycorrhiza within the root system of species of Inga to prevent the rain from washing phosphorus out of the soil.

In some more complex relationships, mycorrhizal fungi do not just collect immobilised soil nutrients, but connect individual plants together by mycorrhizal networks that transport water, carbon, and other nutrients directly from plant to plant through underground hyphal networks.

Suillus tomentosus, a basidiomycete fungus, produces specialized structures known as tuberculate ectomycorrhizae with its plant host lodgepole pine (Pinus contorta var. latifolia). These structures have been shown to host nitrogen fixing bacteria which contribute a significant amount of nitrogen and allow the pines to colonize nutrient-poor sites.

Mechanisms

The mechanisms by which mycorrhizae increase absorption include some that are physical and some that are chemical. Physically, most mycorrhizal mycelia are much smaller in diameter than the smallest root or root hair, and thus can explore soil material that roots and root hairs cannot reach, and provide a larger surface area for absorption. Chemically, the cell membrane chemistry of fungi differs from that of plants. For example, they may secrete organic acids that dissolve or chelate many ions, or release them from minerals by ion exchange. Mycorrhizae are especially beneficial for the plant partner in nutrient-poor soils.

Disease, drought and salinity resistance and its correlation to mycorrhizae

Mycorrhizal plants are often more resistant to diseases, such as those caused by microbial soil-borne pathogens. These associations have been found to assist in plant defense both above and belowground. Mycorrhizas have been found to excrete enzymes that are toxic to soil borne organisms such as nematodes. More recent studies have shown that mycorrhizal associations result in a priming effect of plants that essentially acts as a primary immune response. When this association is formed a defense response is activated similarly to the response that occurs when the plant is under attack. As a result of this inoculation, defense responses are stronger in plants with mycorrhizal associations.

AMF was also significantly correlated with soil biological fertility variables such as soil fungi and soil bacteria, including soil disease. Furthermore, AMF was significantly correlated with soil physical variable, but only with water level and not with aggregate stability. and are also more resistant to the effects of drought. The significance of arbuscular mycorrhizal fungi includes alleviation of salt stress and its beneficial effects on plant growth and productivity. Although salinity can negatively affect arbuscular mycorrhizal fungi, many reports show improved growth and performance of mycorrhizal plants under salt stress conditions 

Resistance to insects

Recent research has shown that plants connected by mycorrihzal fungi can use these underground connections to produce and receive warning signals. Specifically, when a host plant is attacked by an aphid, the plant signals surrounding connected plants of its condition. The host plant releases volatile organic compounds (VOCs) that attract the insect's predators. The plants connected by mycorrhizal fungi are also prompted to produce identical VOCs that protect the uninfected plants from being targeted by the insect. Additionally, this assists the mycorrhizal fungi by preventing the plant’s carbon relocation which negatively affects the fungi’s growth and occurs when the plant is attacked by herbivores.

Colonization of barren soil

Plants grown in sterile soils and growth media often perform poorly without the addition of spores or hyphae of mycorrhizal fungi to colonise the plant roots and aid in the uptake of soil mineral nutrients. The absence of mycorrhizal fungi can also slow plant growth in early succession or on degraded landscapes. The introduction of alien mycorrhizal plants to nutrient-deficient ecosystems puts indigenous non-mycorrhizal plants at a competitive disadvantage. This aptitude to colonize barren soil is defined by the category Oligotroph.

Resistance to toxicity

Fungi have been found to have a protective role for plants rooted in soils with high metal concentrations, such as acidic and contaminated soils. Pine trees inoculated with Pisolithus tinctorius planted in several contaminated sites displayed high tolerance to the prevailing contaminant, survivorship and growth. One study discovered the existence of Suillus luteus strains with varying tolerance of zinc. Another study discovered that zinc-tolerant strains of Suillus bovinus conferred resistance to plants of Pinus sylvestris. This was probably due to binding of the metal to the extramatricial mycelium of the fungus, without affecting the exchange of beneficial substances.

Occurrence of mycorrhizal associations

At around 400 million years old, the Rhynie chert contains an assemblage of fossil plants preserved in sufficient detail that mycorrhizas have been observed in the stems of Aglaophyton major.

Mycorrhizas are present in 92% of plant families studied (80% of species), with arbuscular mycorrhizas being the ancestral and predominant form, and the most prevalent symbiotic association found in the plant kingdom. The structure of arbuscular mycorrhizas has been highly conserved since their first appearance in the fossil record, with both the development of ectomycorrhizas, and the loss of mycorrhizas, evolving convergently on multiple occasions.

Discovery

Associations of fungi with the roots of plants have been known since at least the mid-19th century. However early observers simply recorded the fact without investigating the relationships between the two organisms. This symbiosis was studied and described by Franciszek Kamieński in 1879–1882. Further research was carried out by Albert Bernhard Frank, who introduced the term mycorrhiza in 1885.

Climate change and ecosystems

From Wikipedia, the free encyclopedia

Rainforest ecosystems are rich in biodiversity. This is the Gambia River in Senegal's Niokolo-Koba National Park.

 

This article is about climate change and ecosystems. Future climate change is expected to affect particular ecosystems, including tundra, mangroves, coral reefs, and caves.

General

Unchecked global warming could affect most terrestrial ecoregions. Increasing global temperature means that ecosystems will change; some species are being forced out of their habitats (possibly to extinction) because of changing conditions, while others are flourishing. Secondary effects of global warming, such as lessened snow cover, rising sea levels, and weather changes, may influence not only human activities but also the ecosystem. 

For the IPCC Fourth Assessment Report, experts assessed the literature on the impacts of climate change on ecosystems. Rosenzweig et al. (2007) concluded that over the last three decades, human-induced warming had likely had a discernible influence on many physical and biological systems (p. 81). Schneider et al. (2007) concluded, with very high confidence, that regional temperature trends had already affected species and ecosystems around the world (p. 792). With high confidence, they concluded that climate change would result in the extinction of many species and a reduction in the diversity of ecosystems (p. 792).

• Terrestrial ecosystems and biodiversity: With a warming of 3 °C, relative to 1990 levels, it is likely that global terrestrial vegetation would become a net source of carbon (Schneider et al., 2007:792). With high confidence, Schneider et al. (2007:788) concluded that a global mean temperature increase of around 4 °C (above the 1990-2000 level) by 2100 would lead to major extinctions around the globe.
• Marine ecosystems and biodiversity: With very high confidence, Schneider et al. (2007:792) concluded that a warming of 2 °C above 1990 levels would result in mass mortality of coral reefs globally. In addition, several studies dealing with planktonic organisms and modelling have shown that temperature plays a transcendental role in marine microbial food webs, which may have a deep influence on the biological carbon pump of marine planktonic pelagic and mesopelagic ecosystems.
• Freshwater ecosystems: Above about a 4 °C increase in global mean temperature by 2100 (relative to 1990-2000), Schneider et al. (2007:789) concluded, with high confidence, that many freshwater species would become extinct.

Impacts

Studying the association between Earth climate and extinctions over the past 520 million years, scientists from the University of York write, "The global temperatures predicted for the coming centuries may trigger a new ‘mass extinction event’, where over 50 per cent of animal and plant species would be wiped out."

Many of the species at risk are Arctic and Antarctic fauna such as polar bears and emperor penguins. In the Arctic, the waters of Hudson Bay are ice-free for three weeks longer than they were thirty years ago, affecting polar bears, which prefer to hunt on sea ice. Species that rely on cold weather conditions such as gyrfalcons, and snowy owls that prey on lemmings that use the cold winter to their advantage may be hit hard. Marine invertebrates enjoy peak growth at the temperatures they have adapted to, regardless of how cold these may be, and cold-blooded animals found at greater latitudes and altitudes generally grow faster to compensate for the short growing season. Warmer-than-ideal conditions result in higher metabolism and consequent reductions in body size despite increased foraging, which in turn elevates the risk of predation. Indeed, even a slight increase in temperature during development impairs growth efficiency and survival rate in rainbow trout.

Rising temperatures are beginning to have a noticeable impact on birds, and butterflies have shifted their ranges northward by 200 km in Europe and North America. Plants lag behind, and larger animals' migration is slowed down by cities and roads. In Britain, spring butterflies are appearing an average of 6 days earlier than two decades ago.

A 2002 article in Nature surveyed the scientific literature to find recent changes in range or seasonal behaviour by plant and animal species. Of species showing recent change, 4 out of 5 shifted their ranges towards the poles or higher altitudes, creating "refugee species". Frogs were breeding, flowers blossoming and birds migrating an average 2.3 days earlier each decade; butterflies, birds and plants moving towards the poles by 6.1 km per decade. A 2005 study concludes human activity is the cause of the temperature rise and resultant changing species behaviour, and links these effects with the predictions of climate models to provide validation for them. Scientists have observed that Antarctic hair grass is colonizing areas of Antarctica where previously their survival range was limited.

Mechanistic studies have documented extinctions due to recent climate change: McLaughlin et al. documented two populations of Bay checkerspot butterfly being threatened by precipitation change. Parmesan states, "Few studies have been conducted at a scale that encompasses an entire species" and McLaughlin et al. agreed "few mechanistic studies have linked extinctions to recent climate change." Daniel Botkin and other authors in one study believe that projected rates of extinction are overestimated. For "recent" extinctions, see Holocene extinction. 

Many species of freshwater and saltwater plants and animals are dependent on glacier-fed waters to ensure a cold water habitat that they have adapted to. Some species of freshwater fish need cold water to survive and to reproduce, and this is especially true with salmon and cutthroat trout. Reduced glacier runoff can lead to insufficient stream flow to allow these species to thrive. Ocean krill, a cornerstone species, prefer cold water and are the primary food source for aquatic mammals such as the blue whale. Alterations to the ocean currents, due to increased freshwater inputs from glacier melt, and the potential alterations to thermohaline circulation of the worlds oceans, may affect existing fisheries upon which humans depend as well. 

The white lemuroid possum, only found in the Daintree mountain forests of northern Queensland, may be the first mammal species to be driven extinct by global warming in Australia. In 2008, the white possum has not been seen in over three years. The possums cannot survive extended temperatures over 30 °C (86 °F), which occurred in 2005.

A 27-year study of the largest colony of Magellanic penguins in the world, published in 2014, found that extreme weather caused by climate change is responsible for killing 7% of penguin chicks per year on average, and in some years studied climate change accounted for up to 50% of all chick deaths. Since 1987, the number of breeding pairs in the colony has reduced by 24%.

Climate change is leading to a mismatch between the snow camouflage of arctic animals such as snowshoe hares with the increasingly snow-free landscape.

Forests

Change in Photosynthetic Activity in Northern Forests 1982-2003; NASA Earth Observatory

 

Pine forests in British Columbia have been devastated by a pine beetle infestation, which has expanded unhindered since 1998 at least in part due to the lack of severe winters since that time; a few days of extreme cold kill most mountain pine beetles and have kept outbreaks in the past naturally contained. The infestation, which (by November 2008) has killed about half of the province's lodgepole pines (33 million acres or 135,000 km²) is an order of magnitude larger than any previously recorded outbreak. One reason for unprecedented host tree mortality may be due to that the mountain pine beetles have higher reproductive success in lodgepole pine trees growing in areas where the trees have not experienced frequent beetle epidemics, which includes much of the current outbreak area. In 2007 the outbreak spread, via unusually strong winds, over the continental divide to Alberta. An epidemic also started, be it at a lower rate, in 1999 in Colorado, Wyoming, and Montana. The United States forest service predicts that between 2011 and 2013 virtually all 5 million acres (20,000 km²) of Colorado’s lodgepole pine trees over five inches (127 mm) in diameter will be lost.

As the northern forests are a carbon sink, while dead forests are a major carbon source, the loss of such large areas of forest has a positive feedback on global warming. In the worst years, the carbon emission due to beetle infestation of forests in British Columbia alone approaches that of an average year of forest fires in all of Canada or five years worth of emissions from that country's transportation sources.

Besides the immediate ecological and economic impact, the huge dead forests provide a fire risk. Even many healthy forests appear to face an increased risk of forest fires because of warming climates. The 10-year average of boreal forest burned in North America, after several decades of around 10,000 km² (2.5 million acres), has increased steadily since 1970 to more than 28,000 km² (7 million acres) annually. Though this change may be due in part to changes in forest management practices, in the western U.S., since 1986, longer, warmer summers have resulted in a fourfold increase of major wildfires and a sixfold increase in the area of forest burned, compared to the period from 1970 to 1986. A similar increase in wildfire activity has been reported in Canada from 1920 to 1999.

Forest fires in Indonesia have dramatically increased since 1997 as well. These fires are often actively started to clear forest for agriculture. They can set fire to the large peat bogs in the region and the CO₂released by these peat bog fires has been estimated, in an average year, to be 15% of the quantity of CO₂produced by fossil fuel combustion.

A 2018 study found that trees grow faster due to increased carbon dioxide levels, however, the trees are also eight to twelve percent lighter and denser since 1900. The authors note, "Even though a greater volume of wood is being produced today, it now contains less material than just a few decades ago."

Mountains

Mountains cover approximately 25 percent of earth's surface and provide a home to more than one-tenth of global human population. Changes in global climate pose a number of potential risks to mountain habitats. Researchers expect that over time, climate change will affect mountain and lowland ecosystems, the frequency and intensity of forest fires, the diversity of wildlife, and the distribution of fresh water. 

Studies suggest a warmer climate in the United States would cause lower-elevation habitats to expand into the higher alpine zone. Such a shift would encroach on the rare alpine meadows and other high-altitude habitats. High-elevation plants and animals have limited space available for new habitat as they move higher on the mountains in order to adapt to long-term changes in regional climate. 

Changes in climate will also affect the depth of the mountains snowpacks and glaciers. Any changes in their seasonal melting can have powerful impacts on areas that rely on freshwater runoff from mountains. Rising temperature may cause snow to melt earlier and faster in the spring and shift the timing and distribution of runoff. These changes could affect the availability of freshwater for natural systems and human uses.

Oceans

Ocean Acidification

Estimated annual mean sea surface anthropogenic dissolved inorganic carbon concentration for the present day (normalised to year 2002) from the Global Ocean Data Analysis Project v2 (GLODAPv2) climatology.

 

Annual mean sea surface dissolved oxygen from the World Ocean Atlas 2009. Dissolved oxygen here is in mol O₂m^-3.

 

Ocean acidification poses a severe threat to the earth's natural process of regulating atmospheric C0₂ levels, causing a decrease in water's ability to dissolve oxygen and created oxygen-vacant bodies of water called "dead zones." The ocean absorbs up to 55% of atmospheric carbon dioxide, lessoning the effects of climate change. This diffusion of carbon dioxide into seawater results in three acidic molecules: bicarbonate ion (HCO₃-), aqueous carbon dioxide (CO₂aq), and carbonic acid (H₂CO₃). These three compounds increase the ocean's acidity, decreasing its ph by up to 0.1 per 100ppm (part per million) of atmospheric CO₂. The increase of ocean acidity also decelerates the rate of calcification in salt water, leading to slower growing reefs which support a whopping 25% of marine life. As seen with the great barrier reef, the increase in ocean acidity in not only killing the coral, but also the wildly diverse population of marine inhabitants which coral reefs support.

Dissolved Oxygen

Another issue faced by increasing global temperatures is the decrease of the ocean's ability to dissolve oxygen, one with potentially more severe consequences than other repercussions of global warming. Ocean depths between 100 meters and 1,000 meters are known as "oceanic mid zones" and host a plethora of biologically diverse species, one of which being zooplankton. Zooplankton feed on smaller organisms such as phytoplankton, which are an integral part of the marine food web. Phytoplankton perform photosynthesis, receiving energy from light, and provide sustenance and energy for the larger zooplankton, which provide sustenance and energy for the even larger fish, and so on up the food chain. The increase in oceanic temperatures lowers the ocean's ability to retain oxygen generated from phytoplankton, and therefore reduces the amount of bioavailable oxygen that fish and other various marine wildlife rely on for their survival. This creates marine dead zones, and the phenomenon has already generated multiple marine dead zones around the world, as marine currents effectively "trap" the deoxygenated water.

Combined Impact

Eventually the planet will warm to such a degree that the ocean's ability to dissolve water will no longer exist, resulting in a worldwide dead zone. Dead zones, in combination with ocean acidification, will usher in an era where marine life in most forms will cease to exist, causing a sharp decline in the amount of oxygen generated through bio carbon sequestration, perpetuating the cycle. This disruption to the food chain will cascade upward, thinning out populations of primary consumers, secondary consumers, tertiary consumers, etc., as primary consumers being the initial victims of these phenomenon.

Fresh Water

Disruption to Water-Cycle

The Water Cycle

 

Fresh water covers only 0.8% of the Earth's surface, but contains up to 6% of all life on the planet. However, the impacts climate change deal to its ecosystems are often overlooked. Very few studies showcase the potential results of climate change on large-scale ecosystems which are reliant on freshwater, such as river ecosystems, lake ecosystems, desert ecosystems, etc. However, a comprehensive study published in 2009 delves into the effects to be felt by lotic (flowing) and lentic (still) freshwater ecosystems in the American Northeast. According to the study, persistent rainfall, typically felt year round, will begin to diminish and rates of evaporation will increase, resulting in drier summers and more sporadic periods of precipitation throughout the year. Additionally, a decrease in snowfall is expected, which leads to less runoff in the spring when snow thaws and enters the watershed, resulting in lower-flowing fresh water rivers. This decrease in snowfall also leads to increased runoff during winter months, as rainfall cannot permeate the frozen ground usually covered by water-absorbing snow.  These effects on the water cycle will wreak havoc for indigenous species residing in fresh water lakes and streams.

Salt Water Contamination and Cool Water Species

Eagle River in central Alaska, home to various indigenous freshwater species.

 

Species of fish living in cold or cool water can see a reduction in population of up to 50% in the majority of U.S. fresh water streams, according to most climate change models. The increase in metabolic demands due to higher water temperatures, in combination with decreasing amounts of food will be the main contributors to their decline. Additionally, many fish species (such as salmon) utilize seasonal water levels of streams as a means of reproducing, typically breeding when water flow is high and migrating to the ocean after spawning. Because snowfall is expected to be reduced due to climate change, water runoff is expected to decrease which leads to lower flowing streams, effecting the spawning of millions of salmon. To add to this, rising seas will begin to flood coastal river systems, converting them from fresh water habitats to saline environments where indigenous species will likely perish. In southeast Alaska, the sea rises by 3.96cm/year, redepositing sediment in various river channels and bringing salt water inland. This rise in sea level not only contaminates streams and rivers with saline water, but also the reservoirs they are connected to, where species such as Sockeye Salmon live. Although this species of Salmon can survive in both salt and fresh water, the loss of a body of fresh water stops them from reproducing in the spring, as the spawning process requires fresh water. Undoubtedly, the loss of fresh water systems of lakes and rivers in Alaska will result in the imminent demise of the state's once-abundant population of salmon.

Combined Impact

In general, as the planet warms, the amount of fresh water bodies across the planet decreases, as evaporation rates increase, rain patterns become more sporadic , and watershed patterns become fragmented, resulting in less cyclical water flow in river and stream systems. This disruption to fresh water cycles disrupts the feeding, mating, and migration patterns of organisms reliant on fresh water ecosystems. Additionally, the encroachment of saline water into fresh water river systems endangers indigenous species which can only survive in fresh water.

Ecological productivity

• According to a paper by Smith and Hitz (2003:66), it is reasonable to assume that the relationship between increased global mean temperature and ecosystem productivity is parabolic. Higher carbon dioxide concentrations will favourably affect plant growth and demand for water. Higher temperatures could initially be favourable for plant growth. Eventually, increased growth would peak then decline.
• According to IPCC (2007:11), a global average temperature increase exceeding 1.5–2.5 °C (relative to the period 1980–99), would likely have a predominantly negative impact on ecosystem goods and services, e.g., water and food supply.
• Research done by the Swiss Canopy Crane Project suggests that slow-growing trees only are stimulated in growth for a short period under higher CO₂ levels, while faster growing plants like liana benefit in the long term. In general, but especially in rainforests, this means that liana become the prevalent species; and because they decompose much faster than trees their carbon content is more quickly returned to the atmosphere. Slow growing trees incorporate atmospheric carbon for decades.

Species migration

In 2010, a gray whale was found in the Mediterranean Sea, even though the species had not been seen in the North Atlantic Ocean since the 18th century. The whale is thought to have migrated from the Pacific Ocean via the Arctic. Climate Change & European Marine Ecosystem Research (CLAMER) has also reported that the Neodenticula seminae alga has been found in the North Atlantic, where it had gone extinct nearly 800,000 years ago. The alga has drifted from the Pacific Ocean through the Arctic, following the reduction in polar ice.

In the Siberian subarctic, species migration is contributing to another warming albedo-feedback, as needle-shedding larch trees are being replaced with dark-foliage evergreen conifers which can absorb some of the solar radiation that previously reflected off the snowpack beneath the forest canopy. It has been projected many fish species will migrate towards the North and South poles as a result of climate change, and that many species of fish near the Equator will go extinct as a result of global warming.

Migratory birds are especially at risk for endangerment due to the extreme dependability on temperature and air pressure for migration, foraging, growth, and reproduction. Much research has been done on the effects of climate change on birds, both for future predictions and for conservation. The species said to be most at risk for endangerment or extinction are populations that are not of conservation concern. It is predicted that a 3.5 degree increase in surface temperature will occur by year 2100, which could result in between 600 and 900 extinctions, which mainly will occur in the tropical environments.

Agriculture

Droughts have been occurring more frequently because of global warming and they are expected to become more frequent and intense in Africa, southern Europe, the Middle East, most of the Americas, Australia, and Southeast Asia. Their impacts are aggravated because of increased water demand, population growth, urban expansion, and environmental protection efforts in many areas. Droughts result in crop failures and the loss of pasture grazing land for livestock.

 

Droughts are becoming more frequent and intense in arid and semiarid western North America as temperatures have been rising, advancing the timing and magnitude of spring snow melt floods and reducing river flow volume in summer. Direct effects of climate change include increased heat and water stress, altered crop phenology, and disrupted symbiotic interactions. These effects may be exacerbated by climate changes in river flow, and the combined effects are likely to reduce the abundance of native trees in favor of non-native herbaceous and drought-tolerant competitors, reduce the habitat quality for many native animals, and slow litter decomposition and nutrient cycling. Climate change effects on human water demand and irrigation may intensify these effects. By 2012, North American corn prices had risen to a record $8.34 per bushel in August, leaving 20 of the 211 U.S. ethanol fuel plants idle.

Tipping points in the climate system

From Wikipedia, the free encyclopedia

Possible tipping elements in the climate system.

A tipping point in the climate system is a threshold that, when exceeded, can lead to large changes in the state of the system. Potential tipping points have been identified in the physical climate system, in impacted ecosystems, and sometimes in both. For instance, feedback from the global carbon cycle is a driver for the transition between glacial and interglacial periods, with orbital forcing providing the initial trigger. Earth's geologic temperature record includes many more examples of geologically rapid transitions between different climate states.

Climate tipping points are of particular interest in reference to concerns about climate change in the modern era. Possible tipping point behaviour has been identified for the global mean surface temperature by studying self-reinforcing feedbacks and the past behavior of Earth's climate system. Self-reinforcing feedbacks in the carbon cycle and planetary reflectivity could trigger a cascading set of tipping points that lead the world into a hothouse climate state.

Large-scale components of the Earth system that may pass a tipping point have been referred to as tipping elements. Tipping elements are found in the Greenland and Antarctic ice sheets, possibly causing tens of meters of sea level rise. These tipping points are not always abrupt. For example, at some level of temperature rise the melt of a large part of the Greenland ice sheet and/or West Antarctic Ice Sheet will become inevitable; but the ice sheet itself may persist for many centuries. Some tipping elements, like the collapse of ecosystems, are irreversible.

Definition

The IPCC AR5 defines a tipping point as an irreversible change in the climate system. It states that the precise levels of climate change sufficient to trigger a tipping point remain uncertain, but that the risk associated with crossing multiple tipping points increases with rising temperature. A more broad definition of tipping points is sometimes used as well, which includes abrupt but reversible tipping points.

Tipping point behaviour in the climate can also be described in mathematical terms. Tipping points are then seen as any type of bifurcation with hysteresis. Hysteresis is the dependence of the state of a system on its history. For instance, depending on how warm and cold it was in the past, there can be differing amounts of ice present on the poles at the same concentration of greenhouse gases or temperature.

In the context of climate change, an "adaptation tipping point" has been defined as "the threshold value or specific boundary condition where ecological, technical, economic, spatial or socially acceptable limits are exceeded."

Tipping points for global temperature

There are many positive and negative feedbacks to global temperatures and the carbon cycle that have been identified. The IPCC reports that feedbacks to increased temperatures are net positive for the remainder of this century, with the impact of cloud cover the largest uncertainty. IPCC carbon cycle models show higher ocean uptake of carbon corresponding to higher concentration pathways, but land carbon uptake is uncertain due to the combined effect of climate change and land use changes.

The geologic record of temperature and greenhouse gas concentration allows climate scientists to gather information on climate feedbacks that lead to different climate states, such as the Late Quaternary (past 1.2 million years), the Pliocene period five million years ago and the Cretaceous period, 100 million years ago. Combining this information with the understanding of current climate change resulted in the finding that "A 2 °C warming could activate important tipping elements, raising the temperature further to activate other tipping elements in a domino-like cascade that could take the Earth System to even higher temperatures".

The speed of tipping point feedbacks is a critical concern and the geologic record often fails to provide clarity as to whether past temperature changes have taken only a few decades or many millennia of time. For instance, a tipping point that was once feared to be abrupt and overwhelming is the release of clathrate compounds buried in seabeds and seabed permafrost, but that feedback is now thought to be chronic and long term.

Some individual feedbacks may be strong enough to trigger tipping points on their own. A 2019 study predicts that if greenhouse gases reach three times the current level of atmospheric carbon dioxide that stratocumulus clouds could abruptly disperse, contributing an additional 8 degrees Celsius of warming.

Runaway greenhouse effect

The runaway greenhouse effect is used in astronomical circles to refer to a greenhouse effect that is so extreme that oceans boil away and render a planet uninhabitable, an irreversible climate state that happened on Venus. The IPCC Fifth Assessment Report states that "a 'runaway greenhouse effect'—analogous to Venus—appears to have virtually no chance of being induced by anthropogenic activities." Venus-like conditions on the Earth require a large long-term forcing that is unlikely to occur until the sun brightens by a few tens of percents, which will take a few billion years.

While a runaway greenhouse effect on Earth is virtually impossible, there are indications that Earth could enter a moist greenhouse state that renders large parts of Earth uninhabitable if the climate forcing is large enough to make water vapour (H₂O) a major atmospheric constituent. Conceivable levels of human-made climate forcing would increase water vapour to about 1% of the atmosphere's mass, thus increasing the rate of hydrogen escape to space. If such a forcing were entirely due to CO₂, the weathering process would remove the excess atmospheric CO₂ well before the ocean was significantly depleted.

Tipping elements

Large scale tipping elements

A smooth or abrupt change in temperature can trigger global-scale tipping points. In the cryosphere these include the irreversible melting of Greenland and Antarctic ice sheets. In Greenland, a positive feedback cycle exists between melting and surface elevation. At lower elevations, temperatures are higher, leading to additional melting. This feedback loop can become so strong that irreversible melting occurs. Marine ice sheet instability could trigger a tipping point in West Antarctica. Crossing either of these tipping points leads to accelerated global sea level rise.

When fresh water gets released as a consequence of Greenland melting, a threshold may be crossed which leads to disruption of the thermohaline circulation. The thermohaline circulation transports heat northward which is important for temperature regulation in the Atlantic region. Risks for a complete shutdown are low to moderate under the Paris agreement levels of warming.

Other examples of possible large scale tipping elements are a shift in El Niño–Southern Oscillation. After crossing a tipping point, the warm phase (El Niño) would start to occur more often. Lastly, the southern ocean, which now absorbs a lot of carbon, might switch to a state where it does not do this anymore.

Regional tipping elements

Climate change can trigger regional tipping points as well. Examples are the disappearance of Arctic sea ice, the establishment of woody species in tundra, permafrost loss, the collapse of the monsoon of South Asia and a strengthening of the West African monsoon which would lead to greening of the Sahara and Sahel. Deforestation may trigger a tipping point in rainforests. As rain forests recycle a large part of their rainfall, when a portion of the forest is destroyed local droughts may threaten the remainder. Finally, boreal forests are considered a tipping element as well. Local warming causes trees to die at a higher rate than before, in proportion to the rise in temperature. As more trees die, the woodland becomes more open, leading to further warming and making forests more susceptible to fire. The tipping point is difficult to predict, but is estimated to be between 3–4 °C of global temperature rise.

Cascading tipping points

Crossing a threshold in one part of the climate system may trigger another tipping element to tip into a new state. These are so-called cascading tipping points. Ice loss in West Antarctica and Greenland will significantly alter ocean circulation. Sustained warming of the northern high latitudes as a result of this process could activate tipping elements in that region, such as permafrost degradation, loss of Arctic sea ice, and Boreal forest dieback. This illustrates that even at relatively low levels of global warming, relatively stable tipping elements may be activated.

Early warning signals

For some of the tipping points described above, it may be possible to detect whether that part of the climate system is getting closer to a tipping point. All parts of the climate system are sometimes disturbed by weather events. After the disruption, the system moves back to its equilibrium. A storm may damage sea ice, which grows back after the storm has passed. If a system is getting closer to tipping, this restoration to its normal state might take increasingly longer, which can be used as a warning sign of tipping.

Tipping point effects

If the climate tips into a hothouse Earth scenario, some scientists warn of food and water shortages, hundreds of millions of people being displaced by rising sea levels, unhealthy and unlivable conditions, and coastal storms having larger impacts. Runaway climate change of 4–5 °C can make swathes of the planet around the equator uninhabitable, with sea levels up to 60 metres (197 ft) higher than they are today threatening coastal cities. Humans cannot survive if the air is too moist and hot, which would happen for the majority of human populations if global temperatures rise by 11–12 °C, as land masses warm faster than the global average. Effects like these have been popularized in books like The Uninhabitable Earth, which climate change deniers refer to as sensationalized "climate disaster porn".