Root hairs or absorbent hairs, are outgrowths of epidermal cells, specialized cells at the tip of a plant root. They are lateral extensions of a single cell and are only rarely branched. They are found in the region of maturation, of the root. Root hair cells improve plant water absorption by increasing root surface area to volume ratio which allows the root hair cell to take in more water. The large vacuole inside root hair cells makes this intake much more efficient. Root hairs are also important for nutrient uptake as they are main interface between plants and mycorrhizal fungi.
The function of all root hairs is to collect water and mineral nutrients in the soil to be sent throughout the plant. In roots, most water absorption happens through the root hairs. The length of root hairs allows them to penetrate between soil particles and prevents harmful bacterial organisms from entering the plant through the xylem vessels. [1] Increasing the surface area of these hairs makes plants more efficient in absorbing nutrients and interacting with microbes. [2] As root hair cells do not carry out photosynthesis, they do not contain chloroplasts.
Root hairs form an important surface as they are needed to absorb most of the water and nutrients needed for the plant. They are also directly involved in the formation of root nodules in legume plants. The root hairs curl around the bacteria, which allows for the formation of an infection thread into the dividing cortical cells to form the nodule. [3]
Having a large surface area, the active uptake of water and minerals through root hairs is highly efficient. Root hair cells also secrete acids (e.g., malic and citric acid), which solubilize minerals by changing their oxidation state, making the ions easier to absorb. [4]
Root hair cells vary between 15 and 17 micrometers in diameter, and 80 and 1,500 micrometers in length. [5] Root hairs are found only in the zone of maturation, also called the zone of differentiation. [6] They are not found in the zone of elongation, possibly because older root hairs are sheared off as the root elongates and moves through the soil. [7] Root hairs grow quickly, at least 1 μm/min, making them particularly useful for research on cell expansion. [8] Just prior to and during root hair cell development, there is elevated phosphorylase activity. [9]
Root hairs are essential for healthy plant nutrition, especially through their interactions with symbiotic fungi. Symbiotic fungi and root hairs produce mycorrhizal symbioses like arbuscular mycorrhiza, formed by AM fungi, and ectomycorrhiza, formed by EM fungi. [10] These are very common, [11] occurring in 90% of terrestrial plant species, [12] because of the benefits it brings to both the fungus and plant.
Formation of this relationship for EM fungi begins with the colonization of the root hairs. This process begins when the EM fungus adheres to the root hair from the soil. [13] The fungus then secretes diffusible factors, to which root hairs are highly sensitive, allowing the hyphae to penetrate into the epidermal cells and create a Hartig net in the first layers of the root cortex. [13] This highly branched structure serves as an interface between the two organisms as fungal cells adapt to the exchanges that occur between the plant and fungus. [14] This process is similar to how AM fungi colonize root hairs, but instead of diffusible factors, they secrete hydrolases to relax the cell wall, which allows hyphae to enter, and there is no Hartig net. [13]
Various effects of fungal colonization in root hairs show that this relationship is beneficial to both plant and fungal species, but the main effect is on root hair growth. Fungi actually affect the growth of root hairs if there is water or nutrient deficiency. [13] Since both of these organisms require nutrients and water, their cooperation is essential to their mutual survival. Upon detection of deficiency, the drought stress response of the plant is triggered, causing growth of the root hairs. [12] The mycorrhizae of the fungus then uses its extended system to help the plant find the correct area of nutrition, signaling the direction in which the roots should grow. [13] This makes root growth more efficient, preserving energy for other metabolic processes, which in turn benefits the fungus that feeds off those metabolic products.
When a new root hair cell grows, it excretes a hormone that inhibits the growth of root hairs in nearby cells. This ensures equal and efficient distribution of the actual hairs on these cells.[ citation needed ]
Repotting or transplanting a plant can result in root hair cells being pulled off, perhaps to a significant extent, which can cause wilting. [15]
Mycelium is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Its normal form is that of branched, slender, entangled, anastomosing, hyaline threads. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates into a monokaryotic mycelium, which cannot reproduce sexually; when two compatible monokaryotic mycelia join and form a dikaryotic mycelium, that mycelium may form fruiting bodies such as mushrooms. A mycelium may be minute, forming a colony that is too small to see, or may grow to span thousands of acres as in Armillaria.
A mycorrhiza 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, the plant root system and its surroundings. Mycorrhizae play important roles in plant nutrition, soil biology, and soil chemistry.
An arbuscular mycorrhiza (AM) is a type of mycorrhiza in which the symbiont fungus penetrates the cortical cells of the roots of a vascular plant forming arbuscules. Arbuscular mycorrhiza is a type of endomycorrhiza along with ericoid mycorrhiza and orchid mycorrhiza. They are characterized by the formation of unique tree-like structures, the arbuscules. In addition, globular storage structures called vesicles are often encountered.
The rhizosphere is the narrow region of soil or substrate that is directly influenced by root secretions and associated soil microorganisms known as the root microbiome. Soil pores in the rhizosphere can contain many bacteria and other microorganisms that feed on sloughed-off plant cells, termed rhizodeposition, and the proteins and sugars released by roots, termed root exudates. This symbiosis leads to more complex interactions, influencing plant growth and competition for resources. Much of the nutrient cycling and disease suppression by antibiotics required by plants occurs immediately adjacent to roots due to root exudates and metabolic products of symbiotic and pathogenic communities of microorganisms. The rhizosphere also provides space to produce allelochemicals to control neighbours and relatives.
Glomeromycota are one of eight currently recognized divisions within the kingdom Fungi, with approximately 230 described species. Members of the Glomeromycota form arbuscular mycorrhizas (AMs) with the thalli of bryophytes and the roots of vascular land plants. Not all species have been shown to form AMs, and one, Geosiphon pyriformis, is known not to do so. Instead, it forms an endocytobiotic association with Nostoc cyanobacteria. The majority of evidence shows that the Glomeromycota are dependent on land plants for carbon and energy, but there is recent circumstantial evidence that some species may be able to lead an independent existence. The arbuscular mycorrhizal species are terrestrial and widely distributed in soils worldwide where they form symbioses with the roots of the majority of plant species (>80%). They can also be found in wetlands, including salt-marshes, and associated with epiphytic plants.
Glomus aggregatum is an arbuscular mycorrhizal fungus used as a soil inoculant in agriculture and horticulture. Like other species in this phylum it forms obligate symbioses with plant roots, where it obtains carbon (photosynthate) from the host plant in exchange for nutrients and other benefits.
The ericoid mycorrhiza is a mutualistic relationship formed between members of the plant family Ericaceae and several lineages of mycorrhizal fungi. This symbiosis represents an important adaptation to acidic and nutrient poor soils that species in the Ericaceae typically inhabit, including boreal forests, bogs, and heathlands. Molecular clock estimates suggest that the symbiosis originated approximately 140 million years ago.
Microbial inoculants, also known as soil inoculants or bioinoculants, are agricultural amendments that use beneficial rhizosphericic or endophytic microbes to promote plant health. Many of the microbes involved form symbiotic relationships with the target crops where both parties benefit (mutualism). While microbial inoculants are applied to improve plant nutrition, they can also be used to promote plant growth by stimulating plant hormone production. Although bacterial and fungal inoculants are common, inoculation with archaea to promote plant growth is being increasingly studied.
The Hartig net is the network of inward-growing hyphae, that extends into the plant host root, penetrating between plant cells in the root epidermis and cortex in ectomycorrhizal symbiosis. This network is the internal component of fungal morphology in ectomycorrhizal symbiotic structures formed with host plant roots, in addition to a hyphal mantle or sheath on the root surface, and extramatrical mycelium extending from the mantle into the surrounding soil. The Hartig net is the site of mutualistic resource exchange between the fungus and the host plant. Essential nutrients for plant growth are acquired from the soil by exploration and foraging of the extramatrical mycelium, then transported through the hyphal network across the mantle and into the Hartig net, where they are released by the fungi into the root apoplastic space for uptake by the plant. The hyphae in the Hartig net acquire sugars from the plant root, which are transported to the external mycelium to provide a carbon source to sustain fungal growth.
Nitrogen nutrition in the arbuscular mycorrhizal system refers to...
The mycorrhizosphere includes "roots, hyphae of the directly connected mycorrhizal fungi, associated microorganisms and the soil within their direct influence". It is the region in which nutrients released from the root and the fungus increase the microbial population and its activities. The roots of most terrestrial plants, including most crop plants and almost all woody plants, are colonized by mycorrhiza-forming symbiotic fungi. In this relationship, the plant roots are infected by a fungus, but the rest of the fungal mycelium continues to grow through the soil, digesting and absorbing nutrients and water and sharing these with its plant host. The fungus in turn benefits by receiving photosynthetic sugars from its host.
Soil carbon storage is an important function of terrestrial ecosystems. Soil contains more carbon than plants and the atmosphere combined. Understanding what maintains the soil carbon pool is important to understand the current distribution of carbon on Earth, and how it will respond to environmental change. While much research has been done on how plants, free-living microbial decomposers, and soil minerals affect this pool of carbon, it is recently coming to light that mycorrhizal fungi—symbiotic fungi that associate with roots of almost all living plants—may play an important role in maintaining this pool as well. Measurements of plant carbon allocation to mycorrhizal fungi have been estimated to be 5 to 20% of total plant carbon uptake, and in some ecosystems the biomass of mycorrhizal fungi can be comparable to the biomass of fine roots. Recent research has shown that mycorrhizal fungi hold 50 to 70 percent of the total carbon stored in leaf litter and soil on forested islands in Sweden. Turnover of mycorrhizal biomass into the soil carbon pool is thought to be rapid and has been shown in some ecosystems to be the dominant pathway by which living carbon enters the soil carbon pool.
An ectomycorrhiza is a form of symbiotic relationship that occurs between a fungal symbiont, or mycobiont, and the roots of various plant species. The mycobiont is often from the phyla Basidiomycota and Ascomycota, and more rarely from the Zygomycota. Ectomycorrhizas form on the roots of around 2% of plant species, usually woody plants, including species from the birch, dipterocarp, myrtle, beech, willow, pine and rose families. Research on ectomycorrhizas is increasingly important in areas such as ecosystem management and restoration, forestry and agriculture.
Rhizophagus irregularis is an arbuscular mycorrhizal fungus used as a soil inoculant in agriculture and horticulture. Rhizophagus irregularis is also commonly used in scientific studies of the effects of arbuscular mycorrhizal fungi on plant and soil improvement. Until 2001, the species was known and widely marketed as Glomus intraradices, but molecular analysis of ribosomal DNA led to the reclassification of all arbuscular fungi from Zygomycota phylum to the Glomeromycota phylum.
Ectomycorrhizal extramatrical mycelium is the collection of filamentous fungal hyphae emanating from ectomycorrhizas. It may be composed of fine, hydrophilic hypha which branches frequently to explore and exploit the soil matrix or may aggregate to form rhizomorphs; highly differentiated, hydrophobic, enduring, transport structures.
Dark septate endophytes (DSE) are a group of endophytic fungi characterized by their morphology of melanized, septate, hyphae. This group is likely paraphyletic, and contain conidial as well as sterile fungi that colonize roots intracellularly or intercellularly. Very little is known about the number of fungal taxa within this group, but all are in the Ascomycota. They are found in over 600 plant species and across 114 families of angiosperms and gymnosperms and co-occur with other types of mycorrhizal fungi. They have a wide global distribution and can be more abundant in stressed environments. Much of their taxonomy, physiology, and ecology are unknown.
Orchid mycorrhizae are endomycorrhizal fungi which develop symbiotic relationships with the roots and seeds of plants of the family Orchidaceae. Nearly all orchids are myco-heterotrophic at some point in their life cycle. Orchid mycorrhizae are critically important during orchid germination, as an orchid seed has virtually no energy reserve and obtains its carbon from the fungal symbiont.
Mycorrhiza helper bacteria (MHB) are a group of organisms that form symbiotic associations with both ectomycorrhiza and arbuscular mycorrhiza. MHBs are diverse and belong to a wide variety of bacterial phyla including both Gram-negative and Gram-positive bacteria. Some of the most common MHBs observed in studies belong to the genera Pseudomonas and Streptomyces. MHBs have been seen to have extremely specific interactions with their fungal hosts at times, but this specificity is lost with plants. MHBs enhance mycorrhizal function, growth, nutrient uptake to the fungus and plant, improve soil conductance, aid against certain pathogens, and help promote defense mechanisms. These bacteria are naturally present in the soil, and form these complex interactions with fungi as plant root development starts to take shape. The mechanisms through which these interactions take shape are not well-understood and needs further study.
Rhizophagus clarus is an arbuscular mycorrhizal fungus in the family Glomeraceae. The species has been shown to improve nutrient absorption and growth in several agricultural crops but is not typically applied commercially.
The common symbiosis signaling pathway (CSSP) is a signaling cascade in plants that allows them to interact with symbiotic microbes. It corresponds to an ancestral pathway that plants use to interact with arbuscular mycorrhizal fungi (AMF). It is known as "common" because different evolutionary younger symbioses also use this pathway, notably the root nodule symbiosis with nitrogen-fixing rhizobia bacteria. The pathway is activated by both Nod-factor perception, as well as by Myc-factor perception that are released from AMF. The pathway is distinguished from the pathogen recognition pathways, but may have some common receptors involved in both pathogen recognition as well as CSSP. A recent work by Kevin Cope and colleagues showed that ectomycorrhizae also uses CSSP components such as Myc-factor recognition.