Fungi how does it move

Some fungal organisms multiply only asexually, whereas others undergo both asexual reproduction and sexual reproduction with alternation of generations. Most fungi produce a large number of spores, which are haploid cells that can undergo mitosis to form multicellular, haploid individuals.

Like bacteria, fungi play an essential role in ecosystems because they are decomposers and participate in the cycling of nutrients by breaking down organic and inorganic materials to simple molecules. Fungi often interact with other organisms, forming beneficial or mutualistic associations. For example most terrestrial plants form symbiotic relationships with fungi. The roots of the plant connect with the underground parts of the fungus forming mycorrhizae. Through mycorrhizae, the fungus and plant exchange nutrients and water, greatly aiding the survival of both species Alternatively, lichens are an association between a fungus and its photosynthetic partner usually an alga.

Fungi also cause serious infections in plants and animals. For example, Dutch elm disease, which is caused by the fungus Ophiostoma ulmi , is a particularly devastating type of fungal infestation that destroys many native species of elm Ulmus sp.

The elm bark beetle acts as a vector, transmitting the disease from tree to tree. Accidentally introduced in the s, the fungus decimated elm trees across the continent. Many European and Asiatic elms are less susceptible to Dutch elm disease than American elms. In humans, fungal infections are generally considered challenging to treat.

Unlike bacteria, fungi do not respond to traditional antibiotic therapy because they are eukaryotes. Fungal infections may prove deadly for individuals with compromised immune systems.

Fungi have many commercial applications. The food industry uses yeasts in baking, brewing, and cheese and wine making. Many industrial compounds are byproducts of fungal fermentation.

Fungi are the source of many commercial enzymes and antibiotics. Fungi are unicellular or multicellular thick-cell-walled heterotroph decomposers that eat decaying matter and make tangles of filaments. Fungi are eukaryotes and have a complex cellular organization.

As eukaryotes, fungal cells contain a membrane-bound nucleus where the DNA is wrapped around histone proteins. A few types of fungi have structures comparable to bacterial plasmids loops of DNA. Fungal cells also contain mitochondria and a complex system of internal membranes, including the endoplasmic reticulum and Golgi apparatus.

Unlike plant cells, fungal cells do not have chloroplasts or chlorophyll. Many fungi display bright colors arising from other cellular pigments, ranging from red to green to black. The poisonous Amanita muscaria fly agaric is recognizable by its bright red cap with white patches.

Pigments in fungi are associated with the cell wall. They play a protective role against ultraviolet radiation and can be toxic. The poisonous Amanita muscaria : The poisonous Amanita muscaria is native to temperate and boreal regions of North America. The rigid layers of fungal cell walls contain complex polysaccharides called chitin and glucans. Chitin, also found in the exoskeleton of insects, gives structural strength to the cell walls of fungi.

The wall protects the cell from desiccation and predators. Fungi have plasma membranes similar to other eukaryotes, except that the structure is stabilized by ergosterol: a steroid molecule that replaces the cholesterol found in animal cell membranes. Most members of the kingdom Fungi are nonmotile. The vegetative body of a fungus is a unicellular or multicellular thallus. Dimorphic fungi can change from the unicellular to multicellular state depending on environmental conditions.

Unicellular fungi are generally referred to as yeasts. Example of a unicellular fungus : Candida albicans is a yeast cell and the agent of candidiasis and thrush. This organism has a similar morphology to coccus bacteria; however, yeast is a eukaryotic organism note the nucleus. Most fungi are multicellular organisms.

They display two distinct morphological stages: the vegetative and reproductive. The vegetative stage consists of a tangle of slender thread-like structures called hyphae singular, hypha , whereas the reproductive stage can be more conspicuous. The mass of hyphae is a mycelium. It can grow on a surface, in soil or decaying material, in a liquid, or even on living tissue. Example of a mycelium of a fungus : The mycelium of the fungus Neotestudina rosati can be pathogenic to humans.

The fungus enters through a cut or scrape and develops a mycetoma, a chronic subcutaneous infection. Thus, we revisit relevant existing research in fungal ecology with the link between movement and species coexistence in mind. Therefore, this paper has two closely related aims: One aim is to use the movement ecology framework to define the active movement in filamentous fungi and to provide theoretical background to disentangle the ecological function of hyphal and mycelial movements from physiological and developmental functions.

In doing so, we provide a concept that enables movement ecologists to tap into the research in filamentous fungi ecology. Second, we argue that the explicit recognition of active movement and adoption of movement ecology terminology will provide a more comprehensive treatment of the ecological implications of movement in fungi, and will fuel a new line of research in fungal ecology and community assembly.

In sum, we think that our work will benefit both fungal and movement ecology. This type of fungal movement i. For movement ecology, describing filamentous fungi using the common terminology of movement ecology opens an opportunity to challenge the universality of its basic frameworks and terminologies.

We demonstrate this improvement towards greater universality also by showing how other modular organisms, namely clonal plants and slime molds, can fit into our concept. We provide examples of how our concept can aid the comparative ecological studies between these groups, and the use of microbes as model organisms for movement ecology.

As mentioned earlier, fungal dispersal by spores is not within the scope of this article. However, we want to prevent our concept of mycelial active movement from being mis interpreted as an antipode to the seemingly passive dispersal by spores. First, spores can be actively moved by forces generated by the parental mycelium [ 5 , 6 ].

Second, we support the broad definition of navigation and motion capacity sensu Nathan [ 3 ], which accommodates evolved traits such as when or how many spores are released as a part of mycelial navigation capacity; as well as traits which make spores stick to mobile linkers i.

Movement is one of the means by which organisms interact with their environment. It enables them to respond to environmental challenges and to access resources. The first step in the process of adding fungal active movement to the movement ecology framework requires a revision of the definition of movement itself.

We propose the following definition as inclusive for all organisms that interact with their environment in the ways described by the movement ecology framework of Nathan et al.

Based on this definition, we show below how features of fungal morphology and physiology can be described as movement traits, how those traits enable the fungus to respond to its environment, and how these responses affect fungal community assembly. In doing so, we also align the most important movement ecology and fungal biology terminology.

Just like in motile organisms, in filamentous fungi the environmental cues and stimuli can influence the internal state of the filamentous fungus, and steer navigation capacity the translocation of the biomass motion capacity [ 3 ].

This results in a particular spatial location of the fungal biomass in a particular time movement path [ 3 ]. Three different kinds of translocation in hyphae and mycelium can enable a direct response to the environment, and can be recognized as forms of active movement: Hyphal mycelial growth [ 9 , 10 ], transport within the cytoplasm [ 11 , 12 ], and migration retraction of the entire cytoplasm within a hypha [ 13 ]. In motile unitary organisms, the following is realized as three distinct, decoupled processes Fig.

Main movement functions in unitary motile and modular organism filamentous fungus. In motile unitary organism left , the individual interacts with the environment by moving its entire body from one point to another green arrows; a.

Physiological movements orange arrows; c and developmental movement i. A filamentous fungus right has no capacity to move its entire body. If the mycelium at older locations degenerates possibly recycling some of its own biomass while outgrowing to new locations, the summary result can be a change in position of the entire organism, which is very similar to situations in typical motile organisms.

Therefore, also processes such as autophagy should be recognized as movement related traits [ 14 ]. We point out that just like in other actively moving groups, motion capacity in fungi differs radically between species.

For example, Olsson [ 15 ] let different species grow in Petri dishes with a source of concentrated C on one end, and a source of concentrated N on the opposite end, with the gradient of concentrations in between. While some species were able to actively integrate resources across all space and grow in the entire Petri dish, others were only growing in the central part. In our concept, it is pivotal to make a clear distinction between the two main forms of ecologically relevant movements movement capacities in filamentous fungi, i.

For example, the growth of hyphae is of primary interest in the dispersal of bacteria in soil environment, while the cytoplasmic transport acts in clonal subsidizing.

However, it should be noted that in the development of fungal body, cytoplasmic streaming and hyphal growth are closely interrelated. For more details, we refer to the mycofluidics review by Ropert and Seminara [ 6 ].

Both, the growth of hyphae and transport of biomass within the mycelium can be informed , i. In terms of movement ecology, fungi clearly have navigation capacity Fig.

Remarkable navigation capacities are known for example in the grass pathogen Claviceps purpurea the ergot fungus , in which the hypha must pass through several different tissues in order to find its way from the spore germination site to the young floret which it targets [ 18 ]. Movement ecology framework adapted for the biology and ecology of filamentous fungi: Original graphical representation of the framework by Jeltsch et al. Blue boxes are related to fungal active movement enabled by informed growth.

Orange boxes are related to fungal active movement enabled by cytoplasmic transport. When fungal mycelium is grazed, the interplay between the internal state motivation to move and the physiological ability to move , and the navigation capacity can be complex: While intense grazing results unsurprisingly in decreased growth, moderate grazing can trigger reactions which can be either interpreted as compensatory growth, or escape mechanisms.

Hedlund et al. Authors suggest that aerial hyphae which grow in 3D space have a higher chance to escape the grazer within the pores of a natural substrate. In the Fomina et al. Interestingly, the response negative chemotropism decreased with enhanced availability of sucrose in the medium. Translated to movement ecology terms, this study investigated navigation capacity also in the context of the internal state i.

Hyphae of filamentous fungi are able to use chemotaxis to navigate growth towards other hyphae of the same species, for example between mating partners [ 23 ]. Wood decomposing species were also shown to distinguish between different species of competitors, and change their growth between patches accordingly [ 24 ]. For instance, in the Hanson et al. For example, the frequency of branching was increased. This study is also a good example of directional memory in fungi, and its role in navigation.

Perera et al. For example, plant roots are known for releasing chemoattractants, which the hyphae use for navigation [ 20 ]. The extended movement ecology concept predicts that not only does the environment have an effect on the movement path see examples mentioned above , but also the interactions that take place along the movement path have an effect on the environment, influencing community assembly [ 4 ] Fig.

Just like in other groups, in filamentous fungi the focal individual with its particular movement path can act as a mobile link see below for populations of other guilds. At the same time and within the same guild, movement of this individual is an important factor of fungal community assembly affecting intraspecific and interspecific interactions. Below we expand on these two effects of active fungal movement.

The effect of nutrient transport by fungi i. In analogy to the migration of salmon and feeding habits of bears, which result in the creation of nutrient mobile links [ 29 ], also the nutrients transported by hyphae can be accessed and released by mycophagous bacteria [ 30 ].

However, the nutrient links can have a less dramatic form, where the fungus is not destroyed: In nutrient poor and dry microhabitats, populations of bacteria can be maintained by hyphal transport and excretion of nutrients and water [ 31 ]. Mycelia are also able to transport organic contaminants, making them available for biodegradation by soil bacteria [ 32 ]. Hyphae of filamentous fungi also act as genetic mobile linkers for populations of soil bacteria [ 33 , 34 , 35 ], by providing a network of pathways, which bacterial species can use for their dispersal.

In soil, bacteria can typically only move in the water phase. In dry conditions, this can decrease the habitat connectivity. However, connectivity can be improved again by the presence of hyphae surrounded by a water film [ 34 ]. The dispersal ability of bacteria on fungal hyphae appears to be a result of a complicated interplay between the traits of both partners.

Different fungus - bacteria species combinations show different dispersal potential [ 36 , 37 ], and the effect has been already shown to influence bacterial community composition [ 37 ].

Hydrophobicity decreases the dispersal potential of the fungus [ 36 , 38 ]. On the bacterial side, the ability to actively move within the water film is important [ 34 ], although evidence for passive dispersal also exists [ 36 ]. The example of fungal highways and fungal pipelines the terms often used in fungal biology for genetic and resource mobile linkers, respectively also demonstrates how the adoption of general movement ecology concepts in fungal ecology needs to take into account the specifics of microbial communities.

For example, since in bacteria the dispersal propagules are usually metabolically active cells, often the function of genetic and resource mobile linker is closely related. As shown above, the fungus not only serves as a passive scaffold, but the dispersal can be further facilitated by provision of nutrients. Dispersal can be also accompanied by the function of process linkers.

These can be localized pH alterations [ 39 ], or antibiosis : By creating microenvironments with antibacterial properties, fungi can preferentially spread antibiotic resistant strains [ 40 ].

An interesting example is the movement based mutualism between the filamentous fungus Aspergillus fumigatus and the swarming bacterium Paenibacillus vortex in soil. In return, the hyphae of the fungal partner serve as bridges for P. Our knowledge of community interactions and assembly in filamentous fungi is still limited, despite the recent advances in this field [ 1 , 42 , 43 , 44 ]. We argue that this research will benefit from explicit recognition of fungal active movement within its ecological context.

Below we revisit topics related to fungal intraspecific and interspecific interactions through the lens of movement ecology. Namely mycelial outgrowth as a form of dispersal, mycelial and hyphal foraging, interference competition, and mycelial translocation in clonal subsidizing.

Filamentous fungi can regenerate from small hyphal fragments. This means that any type of growth brings also a potential for dispersal. However, this type of dispersal has rarely been addressed in the context of movement ecology, which we think is a missed opportunity. For instance, since colonization is not only restricted to production and release of spores, then addressing the fundamental movement ecology questions of why the fungus grows in exploration mode e.

An example of this approach i. Authors decided to tackle dispersal by mycelial outgrowth and the resource capture foraging as almost synonymous terms. We imagine that this kind of terminology may leave most of the animal ecologists surprised.

However, it follows closely and correctly the biology and movement ecology of filamentous fungi. Fungal ecologists have long recognized the existence of different foraging strategies while a fungus explores resources. These include the creation of mycelial cords dedicated to foraging [ 19 ], ability to cross an obstacle, or decisions to forage in areas with diverging resource supply [ 10 ].

Agerer [ 48 ] describes up to eight foraging strategies that root mutualistic fungi exhibit. Studies of hyphal movements at the microscopic scale also show that foraging strategies space searching algorithms differ between species [ 49 ]. Using the movement ecology framework leads to discussing these findings in terms of coexistence. For example, if species differ in foraging related traits such as the effectiveness of exploring different geometry, then this can lead to spatial niche partitioning.

Interference competition also known as fungal combat is a well-documented factor of fungal community assembly. We believe the movement ecology framework can offer a new perspective here, since active movement plays an important role in two ways: in preemptive competition and in mycelial transport. Preemptive competition has been identified as one of the main drivers of fungal interference competition [ 50 ]. It is known in fungal ecology that the larger the territory of the mycelium at the moment of contact with the competitor, the higher the likelihood of winning the combat [ 50 , 51 , 52 , 53 , 54 ].

The ability of preempting the available space i. Just like with the directed hyphal growth, this type of growth can also be seen as active movement: Because the incentive to translocate biomass across space is not only developmental growth , but also ecological to capture territory and nutrients.

Hence, it is not only an analogy to animal growth increasing the biomass , but also to the animal increasing its fitness by gaining and keeping a territory with its resources biomass translocation. Besides, preemptive growth is also influenced by the navigation capacity of the fungus, and it can be regarded as a trait important in interspecific variability.

In order for the home range advantage territory size to work, the mycelium must not only occupy the resources. Interference competition is resource costly and in an environment with patchy resource distribution the outcomes can depend also on the differences in the ability of the fungus to effectively integrate resources via mycelial transport [ 17 ].

Perhaps because this is an obvious conclusion to make, and because of technical difficulties to measure [ 55 ], mycelial ability to transport has been - to our knowledge - not explicitly taken into account i.

Rather, the size of the territory is usually measured, and the influence of transport on mycelial combat outcome is black-boxed, together with other species traits, such as the ability to produce particular biochemical agents, or morphological fortifications. For example, a recent study by Kolesidis et al. Among them the mycelial extension rate and relative size of the combating mycelia see above: preemptive competition.

The ability to translocate resources is involved in these parameters, and the model parameterized without disentangling it as a separate parameter, can still predict the competition outcome. However - as the authors also discuss - this may not be the case in other instances, for example in a natural environment with patchy resource distribution. Indeed, the experiment was done using a homogenous agar medium.

In nature, where resources are patchily distributed across larger spatial scales, interspecific variability in transport capacity can play a major role. The existence of this kind of variability has been already shown in different contexts than interference competition [ 15 ].

In the context of interference competition, the importance of transport ability, although not assessed as a trait value across species, was directly or indirectly shown in several studies. When extra resource was made available to Hypholoma fasciculare and Phanerochaete velutina , there was no difference in the combative ability related to the position of this resource which was either distal or proximal to the combat zone [ 51 ].

This suggests that the resources from the distal part were readily available in the interference zone. In another study, the resource bases of two species one saprobic and one mycorrhizal were separated by a column of soil. Still, the size of the resource base determined the outcome of the interference interaction.

The size of the resource also determined the morphology at the interference zone. This is an explicit example of the involvement of mycelial translocations in the outcome of interference competition [ 53 ].

Transport can also be hypothesized as one of the reasons behind the observation that spatial configuration of mycelia influences the combat outcome, irrespectively of the mycelia size [ 56 ]. Given our knowledge about the importance of mycelial transport in interference competition, and our knowledge about the variability in transport abilities from different research contexts, we can envision studies which track the impact of mycelial transport on interference competition in a more explicit way: With mycelial translocations being quantified as a movement trait value across various species.

And with the mycelial translocations seen as active movement phenomena, where questions like when and where to move are central. Hence, in a way similar to how movement ecologists look at the relationship between movement and competition.

As described above, filamentous fungi can use mycelial transport to integrate resources from different patches, but this movement ability trait differs among species [ 15 ].

It is therefore possible that species coexistence can be promoted alongside the trade-off between the ability of resource integration and faster growth. Similarly, species may differ in the ability to transport metabolites into the parts of mycelium, where growth is temporarily not possible due to locally adverse environmental conditions for possible trade-off in fungal network cost and transport efficiency, see Heaton et al.

The ability to transport nutrients and metabolites across the entire mycelium i. It connects them for example to clonal plants, where the impact of this form of active movement has been already studied within the coexistence context see below and [ 57 ]. There are several ways in which we expect our concept to facilitate interdisciplinary collaboration among movement ecologists and fungal biologist, which would be beneficial for the development of both fields.

Here, we expand on several specific examples. Our concept can improve the transfer of knowledge from fungal ecology to movement ecology. As shown above, the concept helps translate the relevant knowledge in fungal ecology and biology into a form accessible for movement ecologists.

We argue that this is needed given the research gaps in movement ecology; Holyoak et al. Moreover, they admit that their review was probably biased against microorganismal movement. This happened, because during the screening for relevant articles, they had to exclude several keywords often used in the microorganismal movement research e.

These keywords proved to be impractical, as using them identified a large number of articles relevant rather for the fields of molecular biology and cell biology. In fungal biology, movement phenomena are often described using specific terminology. For example, the results of the studies about fungal space searching algorithms are highly relevant for fungal foraging topics, however the term foraging was not used in the articles which we reviewed [ 26 , 49 , 59 , 60 ].

The main purpose of the fruitbody is to produce spores so that the fungus can spread. Spores of mushrooms form on special hyphae on the surface of thin gills that form in a circle hanging on the underside of the cap.

The cap has a curved shape poroharore so that the rain droplets run off and the spores keep dry. Mushrooms must shed their spores fast as both mushrooms and spores often live for only a few days. If you use a microscope to make the spores look much larger, you can see them clearly. Check out the spore print activity to learn how to make a print from spores of a mushroom.

Using microscopes to identify fungi parts — adapt our Ferns under the microscope activity so students can have a closer look at different fungi — and why not build in some additional learning about How microscopes magnify? Mushroom spores — learn how to make a print from spores of a mushroom. The Science Learning Hub would like to acknowledge Manaaki Whenua — Landcare Research and the writers for their permission and help to adapt this publication for the web.

An electronic version of this teacher guidebook is available to download from Huia Publishers. Add to collection. Activity ideas Using microscopes to identify fungi parts — adapt our Ferns under the microscope activity so students can have a closer look at different fungi — and why not build in some additional learning about How microscopes magnify?

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