R gene

Last updated
Leucine-rich repeat receptor-like protein kinase
Identifiers
SymbolLRRK
Membranome 737
Leucine-rich repeat receptor-like protein
Identifiers
SymbolLRRP
Membranome 605
TIR-NBS-LRR disease resistance proteins
Identifiers
SymbolTIR-NBS-LRR
Membranome 1343
TIR domain plant proteins
Identifiers
SymbolTIRP
Membranome 1344

Resistance genes (R-Genes) are genes in plant genomes that convey plant disease resistance against pathogens by producing R proteins. The main class of R-genes consist of a nucleotide binding domain (NB) and a leucine rich repeat (LRR) domain(s) and are often referred to as (NB-LRR) R-genes or NLRs. [1] Generally, the NB domain binds either ATP/ADP or GTP/GDP. The LRR domain is often involved in protein-protein interactions as well as ligand binding. NB-LRR R-genes can be further subdivided into toll interleukin 1 receptor (TIR-NB-LRR) and coiled-coil (CC-NB-LRR). [2]

Contents

Resistance can be conveyed through a number of mechanisms including:

Once the R protein has detected the presence of a pathogen, the plant can mount a defence against the pathogen. Because R genes confer resistance against specific pathogens, it is possible to transfer an R gene from one plant to another and make a plant resistant to a particular pathogen.

Many plant resistance proteins are single-pass transmembrane proteins that belong to receptor kinases and Toll-like receptors. R genes are of large interest in crop breeding, providing a large part of the immunity required by agricultural pathosystems. [1]

Background

Plant defense mechanisms depend on detection of fungal and bacterial pathogens. R genes protein syntheses are a way of identifying the pathogen effectors and stop their infection throughout the plant system. Molecules essential for pathogen defense are pattern recognition receptors (PRRs), wall associated kinase (WAKs), receptors with nucleotide-binding domain (NLRs) and leucine-rich repeats (LRRs). All these R proteins play roles in detecting and recognizing pathogen effectors, initiating multiple signal transductions inside the plant cell, these signals transductions will lead to different responses that will aid in pathogen destruction and prevention of further infection. These responses are:

Note that plants have various mechanisms to prevent and detect pathogenic infections, but factors such as geography, environment, genetic, and timing can affect the recognition pattern of a pathogen or can have an effect on the recognition of avirulent (avr) pathogens in plants.

Pathogen recognition

R genes synthesize proteins that will aid with the recognition of pathogenic effectors:

Pattern recognition receptors (PRRs)

This receptor is often composed of leucine-rich repeats (LRRs). LRRs have a wide range of bacterial (proteins), fungal (carbohydrates) and virulent (nucleic acids) recognition, this means that LRRs recognizes many different molecules but each LRRs usually has a very specific molecule it detects. The ability of PRRs to recognize various pathogenic components relies on a regulatory protein called brassinosteroid insensitive 1 –associated receptor kinase (BAK1). Once the pathogen has been recognized by PRRs the release of a kinase into the nucleus has been transduced triggering a transcriptional reprogramming.

Wall associated kinase (WAKs)

The plant cell wall is conformed of pectin and other molecules. Pectin has abundant galacturonic acids which is the compound that WAKs recognizes after a foreign invasion in the plant. Every WAKs (WAK1 & WAK2) has an N-terminal which interacts with pectin in the cell wall when pectin is being degraded to galacturonic acids by fungal enzymes.

Pathogen-associated molecular pattern (PAMPs) and damage-associated molecular pattern (DAMPs) are often identified by lectins which is a protein that binds specific carbohydrates.

Nucleotide-binding domain and leucine-rich repeats (NLRs)

Most R genes code for these immune receptor proteins. [1] NLRs shifts its conformation from ADP state to and ATP state which allows it to send as signal transduction. The activation of NLRs is yet to be completely understood, according to current studies suggest that it is subject to multiple regulators (dimerization or oligomerization, epigenetic and transcriptional regulation, alternative splicing, and proteasome-mediated regulation)

Despite all these differences NLRs, PRRs, WAKs, effector trigger immunity (ETI) and PAMP-triggered immunity (PTI) there are certain similarities such as in the mechanism of signal transduction which includes mitogen-protein kinase (MAPK) cascades through phosphorylation which will be, calcium ion signaling.

An overall overview about the mechanical interaction about a plant defense and the ability of a pathogen to infect a plant would be for instance such a common interaction between bacterial flagellin and receptor-like kinase which triggers a basal immunity sending signals through MAP kinase cascades and transcriptional reprogramming mediated by plant WRKY transcription factors (Stephen T). Also plant resistance protein recognize bacterial effectors and programs resistance through ETI responses.

Various other types

The EDS1 family is a family of plant disease resistance proteins including the nominate enhanced disease susceptibility 1/EDS1 itself and phytoalexin deficient 4/PAD4. The best studied examples of EDS1 and PAD4 are the Arabidopsis thaliana § EDS1 family members. [4]

Signal transduction

A plant defense has two different types of immune system, the one that recognizes pathogen/microbes associated molecular patterns (PAMPs), and this is also known as PAMP-triggered immunity (PTI). Plant defense mechanism depends on immune receptors found on the plasma membrane and then the mechanism can sense the pathogen associated molecular patterns (PAMPs) and microbial associated molecular patterns (MAMPs). Detection of PAMPs triggers a physiological change in the cell activated by the pattern recognition receptors (PRRs) initiating a cascade response which through the recognition of PAMPs and MAMPs lead to the plant resistance. The other type of defense is also known as effector-triggered immunity (ETI) which is the second type of defense mediated by R-proteins by detecting photogenic effectors. ETI detects pathogenic factors and initiates a defense response. ETI is a much faster and amplified system than PTI and it develops onto the hypersensitive response (HR) leading the infected host cell to apoptosis. This does not terminate the pathogen cycle, it just slows the cycle down.

Plants have many ways of identifying symbiotic or foreign pathogens; one of these receptors causes fluctuations in the calcium ions and this fluctuation in the calcium ions. A transcription factor plays an important role in defenses against pathogenic invasion.

Pathogen invasion

Despite the sophiscation of plant defenses, some pathogens have evolved ways to overcome these defenses in order to infect and spread.

Pathogen elicitors are molecules that stimulate any plant defense; among these elicitors we can find two types of pathogen derived elicitors, pathogen/microbe associated molecular pattern (PAMPs/MAMPs), and also there is a second type which is produced by plants known as damage or danger associated molecular patterns (DAMPs). PTI is a way of responding against pathogen actions happening outside the cell, but a much stronger response like ETI is generated in response to effectors molecules. Once there is an induced resistance also known as priming, the plant can react faster and stronger to a pathogen attack. A known priming inducer is called β-aminobutyric acid (BABA) which is a non-protein amino acid.

Successful pathogens evolve changes in their chemical conformation in order to avoid detection by PRRs and WAKs.

Some viruses have mechanisms that allow them to avoid or suppress the RNA-mediated defense (RMD) that some viruses induce in non-transgenic plants. Further studies have shown that this suppression of the host defense has been done by HC-protease (HCPro) encoded in the Potyviral genome. It was later stablished that HCPro was a mechanism used to suppress post-transcriptional gene slicing (PTGs). Cucumber mosaic virus (CMV) uses a different protein called 2b (Pfam PF03263) which is also a suppressor of PTGS in Nicotiana benthamiana.

Even though HcPro and the 2b protein have different protein sequence specific to their own virus, both target the same instrument of defense through different mechanisms.

Genetic engineering

R genes are common subjects of gene cloning. Every advance in techniques of sequencing and transfer has eased this process, progressively requiring less linkage drag, expense, and laboratory work over time. In the future even better results are expected from ever larger data sets, across ever larger numbers of individuals and populations, with ever greater resolution due to both more accurate sequencing and post-sequencing computational comparison between individuals. [1] [3]

See also

Related Research Articles

<i>Arabidopsis thaliana</i> Model plant species in the family Brassicaceae

Arabidopsis thaliana, the thale cress, mouse-ear cress or arabidopsis, is a small plant from the mustard family (Brassicaceae), native to Eurasia and Africa. Commonly found along the shoulders of roads and in disturbed land, it is generally considered a weed.

<span class="mw-page-title-main">Flagellin</span> Bacterial protein

Flagellin is a globular protein that arranges itself in a hollow cylinder to form the filament in a bacterial flagellum. It has a mass of about 30,000 to 60,000 daltons. Flagellin is the principal component of bacterial flagella, and is present in large amounts on nearly all flagellated bacteria.

Pathogen-associated molecular patterns (PAMPs) are small molecular motifs conserved within a class of microbes, but not present in the host. They are recognized by toll-like receptors (TLRs) and other pattern recognition receptors (PRRs) in both plants and animals. This allows the innate immune system to recognize pathogens and thus, protect the host from infection.

Pattern recognition receptors (PRRs) play a crucial role in the proper function of the innate immune system. PRRs are germline-encoded host sensors, which detect molecules typical for the pathogens. They are proteins expressed mainly by cells of the innate immune system, such as dendritic cells, macrophages, monocytes, neutrophils, as well as by epithelial cells, to identify two classes of molecules: pathogen-associated molecular patterns (PAMPs), which are associated with microbial pathogens, and damage-associated molecular patterns (DAMPs), which are associated with components of host's cells that are released during cell damage or death. They are also called primitive pattern recognition receptors because they evolved before other parts of the immune system, particularly before adaptive immunity. PRRs also mediate the initiation of antigen-specific adaptive immune response and release of inflammatory cytokines.

<span class="mw-page-title-main">Innate immune system</span> Immunity strategy in living beings

The innate immune system or nonspecific immune system is one of the two main immunity strategies in vertebrates. The innate immune system is an alternate defense strategy and is the dominant immune system response found in plants, fungi, prokaryotes, and invertebrates.

Systemic acquired resistance (SAR) is a "whole-plant" resistance response that occurs following an earlier localized exposure to a pathogen. SAR is analogous to the innate immune system found in animals, and although there are many shared aspects between the two systems, it is thought to be a result of convergent evolution. The systemic acquired resistance response is dependent on the plant hormone, salicylic acid.

<span class="mw-page-title-main">Hypersensitive response</span>

Hypersensitive response (HR) is a mechanism used by plants to prevent the spread of infection by microbial pathogens. HR is characterized by the rapid death of cells in the local region surrounding an infection and it serves to restrict the growth and spread of pathogens to other parts of the plant. It is analogous to the innate immune system found in animals, and commonly precedes a slower systemic response, which ultimately leads to systemic acquired resistance (SAR). HR can be observed in the vast majority of plant species and is induced by a wide range of plant pathogens such as oomycetes, viruses, fungi and even insects.

The gene-for-gene relationship is a concept in plant pathology that plants and their diseases each have single genes that interact with each other during an infection. It was proposed by Harold Henry Flor who was working with rust (Melampsora lini) of flax (Linum usitatissimum). Flor showed that the inheritance of both resistance in the host and parasite ability to cause disease is controlled by pairs of matching genes. One is a plant gene called the resistance (R) gene. The other is a parasite gene called the avirulence (Avr) gene. Plants producing a specific R gene product are resistant towards a pathogen that produces the corresponding Avr gene product. Gene-for-gene relationships are a widespread and very important aspect of plant disease resistance. Another example can be seen with Lactuca serriola versus Bremia lactucae.

<span class="mw-page-title-main">NOD-like receptor</span> Class of proteins

The nucleotide-binding oligomerization domain-like receptors, or NOD-like receptors (NLRs), are intracellular sensors of pathogen-associated molecular patterns (PAMPs) that enter the cell via phagocytosis or pores, and damage-associated molecular patterns (DAMPs) that are associated with cell stress. They are types of pattern recognition receptors (PRRs), and play key roles in the regulation of innate immune response. NLRs can cooperate with toll-like receptors (TLRs) and regulate inflammatory and apoptotic response.

Biotic stress is stress that occurs as a result of damage done to an organism by other living organisms, such as bacteria, viruses, fungi, parasites, beneficial and harmful insects, weeds, and cultivated or native plants. It is different from abiotic stress, which is the negative impact of non-living factors on the organisms such as temperature, sunlight, wind, salinity, flooding and drought. The types of biotic stresses imposed on an organism depend the climate where it lives as well as the species' ability to resist particular stresses. Biotic stress remains a broadly defined term and those who study it face many challenges, such as the greater difficulty in controlling biotic stresses in an experimental context compared to abiotic stress.

<span class="mw-page-title-main">Plant disease resistance</span> Ability of a plant to stand up to trouble

Plant disease resistance protects plants from pathogens in two ways: by pre-formed structures and chemicals, and by infection-induced responses of the immune system. Relative to a susceptible plant, disease resistance is the reduction of pathogen growth on or in the plant, while the term disease tolerance describes plants that exhibit little disease damage despite substantial pathogen levels. Disease outcome is determined by the three-way interaction of the pathogen, the plant and the environmental conditions.

<span class="mw-page-title-main">Effector-triggered immunity</span>

Effector-triggered immunity (ETI) is one of the pathways, along with the pattern-triggered immunity (PTI) pathway, by which the innate immune system recognises pathogenic organisms and elicits a protective immune response. ETI is elicited when an effector protein secreted by a pathogen into the host cell is successfully recognised by the host. Alternatively, effector-triggered susceptibility (ETS) can occur if an effector protein can block the immune response triggered by pattern recognition receptors (PRR) and evade immunity, allowing the pathogen to propagate in the host.

Guard theory is a branch of immunology which concerns the innate sensing of stereotypical consequences of a virulence factor or pathogen. This is in contrast to the classical understanding of recognition by the innate immune system, which involves recognition of distinct microbial structures- pathogen-associated molecular patterns (PAMPs)- with pattern recognition receptors (PRRs). Some of these stereotypical consequences of virulence factors and pathogens may include altered endosomal trafficking and changes in the cytoskeleton. These recognition mechanisms would work to complement classical pattern recognition mechanisms.

Mitogen-activated protein kinase (MAPK) networks are the pathways and signaling of MAPK, which is a protein kinase that consists of amino acids serine and threonine. MAPK pathways have both a positive and negative regulation in plants. A positive regulation of MAPK networks is to help in assisting with stresses from the environment. A negative regulation of MAPK networks is pertaining to a high quantity of reactive oxygen species (ROS) in the plant.

<span class="mw-page-title-main">Leucine-rich repeat receptor like protein kinase</span>

Leucine-rich repeat receptor like protein kinase are plant cell membrane localized Leucine-rich repeat (LRR) receptor kinase that play critical roles in plant innate immunity. Plants have evolved intricate immunity mechanism to combat against pathogen infection by recognizing Pathogen Associated Molecular Patterns (PAMP) and endogenous Damage Associated Molecular Patterns (DAMP). PEPR 1 considered as the first known DAMP receptor of Arabidopsis.

<span class="mw-page-title-main">EF-Tu receptor</span> Pattern-recognition receptor (PRR)

EF-Tu receptor, abbreviated as EFR, is a pattern-recognition receptor (PRR) that binds to the prokaryotic protein EF-Tu in Arabidopsis thaliana. This receptor is an important part of the plant immune system as it allows the plant cells to recognize and bind to EF-Tu, preventing genetic transformation by and protein synthesis in pathogens such as Agrobacterium.

<span class="mw-page-title-main">High Affinity K+ transporter HAK5</span>

High Affinity K+ transporter HAK5 is a transport protein found on the cell surface membrane of plants under conditions of potassium deprivation. It is believed to act as a symporter for protons and the potassium ion, K+. Firstly discovered in barley, receiving the name of HvHAK1, it was soon after identified in the model plant Arabidopsis thaliana and named HAK5. These transporters belongs to the subgroup I of the KT-HAK-KUP family of plant proteins with obvious homology with both bacterial and fungal transport systems, which experienced a major diversification following land conquest. KT-HAK-KUP transporters are one of four different types of K+ transporter within the cell, but are unique as they do not have a putative pore forming domain like the other three; Shaker channels, KCO channels, HKT transporters. It is activated when the plant is situated in low soil with low potassium concentration, and has been shown to be located in higher concentration in the epidermis and vasculature of K+ deprived plants. By turning on, it increases the plants affinity (uptake) of potassium. Potassium plays a vital role in the plants growth, reproduction, immunity, ion homeostasis, and osmosis, which ensures the plants survival. It is the highest cationic molecule within the plant, accounting for 10% of the plants dry weight, which makes its uptake into the plant important. Each plant species has its own HAK5 transporter that is specific to that species and has different levels of affinity to K+. To operate and activate the HAK5 transporter, the external concentration of K+ must be lower than 10μM and up to 200μM. In Arabidopsis plants, when external potassium concentration is lower than 10μM, it is only HAK5 that is involved with the uptake of K+, then between 10 and 200μM both HAK5 and AKT1 are involved with the uptake of K+. HAK5 is coupled with CBL9/CIPK23 kinase's although the mechanism behind this has not yet been understood.

Botrytis–induced kinase 1 (BIK1) is a membrane-anchored enzyme in plants. It is a kinase that provides resistance to necrotrophic and biotrophic pathogens. As its name suggests, BIK1 is only active after being induced by Botrytis infection. When Botrytis cinerea is present, the BIK1 gene is transcribed so that the kinase is present to defend the cell. BIK1 functions to regulate the amount of salicylic acid (SA) present in the cell. When Botrytis cinerea or Alternaria brassicicola or any other necrotrophic pathogen is present, BIK1 is transcribed to regulate the pathogen response mechanisms. When BIK1 is present, SA levels decrease, allowing the nectrotrophic response to take place. When nectrotrophic pathogens are not present, BIK1 is not transcribed and SA levels increase, limiting the necrotrophic resistance pathway. Only the pathogenic defense response that is initiated by BIK1 is dependent on SA levels. Non-pathogenic cellular functions occur independently. In terms of non-pathogenic cellular functions, BIK1 is described as a critical component of ET signaling and PAMP-triggered immunity to pathogens.

PEPR 1 and PEPR2 are homolog kinases that act as enzymes on other proteins. They attach a phosphate group to specific proteins, called phosphorylation. These reactions can cause the function of the phosphorylated proteins to change. Both PEPR 1 and PEPR 2 can be classified as receptor kinases, which serve an important role in immunity in plants. Receptor kinases have the ability to change the conformation of receptors by adding the phosphate group. These specific receptor kinases serve as a pattern recognition receptor, or PRR, that can quickly and efficiently recognize many different molecular patterns or signatures that are unique to each pathogen. They can also detect different danger signals released from the host and respond accordingly. More specifically, the proteins contain leucine-rich repeat segments that interact outside of the cell. This leucine-rich repeat is a structural motif present in some proteins that has specific functions due to its folded structure. This fold can contain many repeating amino acids, but the most common is the hydrophobic leucine, hence the name. PEPR1 and PEPR2 are present in plants and are involved in several immune system processes. Their ability to change the conformation of receptors can have an effect on signaling processes within plants, allowing the plant to have a system of immunity in place in case of an infection or pathogen.

<span class="mw-page-title-main">Fungal effectors</span> Molecules secreted by pathogenic fungi to modulate the hosts immune response

Fungal effectors are proteins or non-proteinaceous molecules secreted by pathogenic fungi into a host organism in order to modulate the host's immune response.

References

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  2. Knepper C, Day B (2010). "From perception to activation: the molecular-genetic and biochemical landscape of disease resistance signaling in plants". The Arabidopsis Book. 8: e012. doi:10.1199/tab.0124. PMC   3244959 . PMID   22303251.
  3. 1 2 Hafeez AN, Arora S, Ghosh S, Gilbert D, Bowden RL, Wulff BB (July 2021). "Creation and judicious application of a wheat resistance gene atlas" (PDF). Molecular Plant. Cell Press. 14 (7): 1053–1070. doi: 10.1016/j.molp.2021.05.014 . PMID   33991673. S2CID   234683221.
  4. Lapin D, Bhandari DD, Parker JE (August 2020). "Origins and Immunity Networking Functions of EDS1 Family Proteins". Annual Review of Phytopathology. Annual Reviews. 58 (1): 253–276. doi:10.1146/annurev-phyto-010820-012840. PMID   32396762. S2CID   218617308.

Further reading