Biotic stress

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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. [1] 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. [2] 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.

Contents

The damage caused by these various living and nonliving agents can appear very similar. [1] Even with close observation, accurate diagnosis can be difficult. [1] For example, browning of leaves on an oak tree caused by drought stress may appear similar to leaf browning caused by oak wilt, a serious vascular disease caused by a fungus, or the browning caused by anthracnose, a fairly minor leaf disease.

Agriculture

Biotic stressors are a major focus of agricultural research, due to the vast economic losses caused to cash crops. The relationship between biotic stress and plant yield affects economic decisions as well as practical development. The impact of biotic injury on crop yield impacts population dynamics, plant-stressor coevolution, and ecosystem nutrient cycling. [3]

Biotic stress also impacts horticultural plant health and natural habitats ecology. It also has dramatic changes in the host recipient. Plants are exposed to many stress factors, such as drought, high salinity or pathogens, which reduce the yield of the cultivated plants or affect the quality of the harvested products. Although there are many kinds of biotic stress, the majority of plant diseases are caused by fungi. [4] Arabidopsis thaliana is often used as a model plant to study the responses of plants to different sources of stress. [5]

In history

Biotic stresses have had huge repercussions for humanity; an example of this is the potato blight, an oomycete which caused widespread famine in England, Ireland and Belgium in the 1840s. [6] Another example is grape phylloxera coming from North America in the 19th century, which led to the Great French Wine Blight. [6]

Today

Losses to pests and disease in crop plants continue to pose a significant threat to agriculture and food security. During the latter half of the 20th century, agriculture became increasingly reliant on synthetic chemical pesticides to provide control of pests and diseases, especially within the intensive farming systems common in the developed world. However, in the 21st century, this reliance on chemical control is becoming unsustainable. Pesticides tend to have a limited lifespan due to the emergence of resistance in the target pests, and are increasingly recognised in many cases to have negative impacts on biodiversity, and on the health of agricultural workers and even consumers. [7]

Tomorrow

Due to the implications of climate change, it is suspected that plants will have increased susceptibility to pathogens. [8] Additionally, elevated threat of abiotic stresses (i.e. drought and heat) are likely to contribute to plant pathogen susceptibility. [8]

Effect on plant growth

Photosynthesis

Many biotic stresses affect photosynthesis, as chewing insects reduce leaf area and virus infections reduce the rate of photosynthesis per leaf area. Vascular-wilt fungi compromise the water transport and photosynthesis by inducing stomatal closure. [6] [9]

Response to stress

Plants have co-evolved with their parasites for several hundred million years. This co-evolutionary process has resulted in the selection of a wide range of plant defences against microbial pathogens and herbivorous pests which act to minimise frequency and impact of attack. These defences include both physical and chemical adaptations, which may either be expressed constitutively, or in many cases, are activated only in response to attack. For example, utilization of high metal ion concentrations derived from the soil allow plants to reduce the harmful effects of biotic stressors (pathogens, herbivores etc.); meanwhile preventing the infliction of severe metal toxicity by way of safeguarding metal ion distribution throughout the plant with protective physiological pathways. [10] Such induced resistance provides a mechanism whereby the costs of defence are avoided until defense is beneficial to the plant. At the same time, successful pests and pathogens have evolved mechanisms to overcome both constitutive and induced resistance in their particular host species. In order to fully understand and manipulate plant biotic stress resistance, we require a detailed knowledge of these interactions at a wide range of scales, from the molecular to the community level. [7]

Inducible defense responses to insect herbivores

In order for a plant to defend itself against biotic stress, it must be able to differentiate between an abiotic and biotic stress. A plants response to herbivores starts with the recognition of certain chemicals that are abundant in the saliva of the herbivores. These compounds that trigger a response in plants are known as elicitors or herbivore-associated molecular patterns (HAMPs). [11] These HAMPs trigger signalling pathways throughout the plant, initiating its defence mechanism and allowing the plant to minimise damage to other regions. These HAMPs trigger signalling pathways throughout the plant, initiating its defence mechanism and allowing the plant to minimise damage to other regions. Phloem feeders, like aphids, do not cause a great deal of mechanical damage to plants, but they are still regarded as pests and can seriously harm crop yields. Plants have developed a defence mechanism using salicylic acid pathway, which is also used in infection stress, when defending itself against phloem feeders. Plants perform a more direct attack on an insects digestive system. The plants do this using proteinase inhibitors. These proteinase inhibitors prevent protein digestion and once in the digestive system of an insect, they bind tightly and specifically to the active site of protein hydrolysing enzymes such as trypsin and chymotrypsin. [11] This mechanism is most likely to have evolved in plants when dealing with insect attack.

Plants detect elicitors in the insects saliva. Once detected, a signal transduction network is activated. The presence of an elicitor causes an influx of Ca2+ ions to be released in to the cytosol. This increase in cytosolic concentration activates target proteins such as Calmodulin and other binding proteins. Downstream targets, such as phosphorylation and transcriptional activation of stimulus specific responses, are turned on by Ca2+ dependent protein kinases. [11] In Arabidopsis, over expression of the IQD1 calmodulin-binding transcriptional regulator leads to inhibitor of herbivore activity. The role of calcium ions in this signal transduction network is therefore important.

Calcium Ions also play a large role in activating a plants defensive response. When fatty acid amides are present in insect saliva, the mitogen-activated protein kinases (MAPKs) are activated. These genes when activated, play a role in the jasmonic acid pathway. [11] The jasmonic acid pathway is also referred to as the Octadecanoid pathway. This pathway is vital for the activation of defence genes in plants. The production of jasmonic acid, a phytohormone, is a result of the pathway. In an experiment using virus-induced gene silencing of two calcium-dependent protein kinases (CDPKs) in a wild tobacco ( Nicotiana attenuata ), it was discovered that the longer herbivory continued the higher the accumulation of jasmonic acid in wild-type plants and in silenced plants, the production of more defence metabolites was seen as well as the decrease in the growth rate of the herbivore used, the tobacco hornworm ( Manduca sexta ). [11] This example demonstrates the importance of MAP kinases in plant defence regulation.

Inducible defense responses to pathogens

Plants are capable of detecting invaders through the recognition of non-self signals despite the lack of a circulatory or immune system like those found in animals. Often a plant's first line of defense against microbes occurs at the plant cell surface and involves the detection of microorganism-associated molecular patterns (MAMPs). [12] MAMPs include nucleic acids common to viruses and endotoxins on bacterial cell membranes which can be detected by specialized pattern-recognition receptors. [13] Another method of detection involves the use of plant immune receptors to detect effector molecules released into plant cells by pathogens. Detection of these signals in infected cells leads to an activation of effector-triggered immunity (ETI), a type of innate immune response. [14]

Both the pattern recognition immunity (PTI) and effector-triggered immunity (ETI) result from the upregulation of multiple defense mechanisms including defensive chemical signaling compounds. [14] An increase in the production of salicylic acid (SA) has been shown to be induced by pathogenic infection. The increase in SA results in the production of pathogenesis related (PR) genes which ultimately increase plant resistance to biotrophic and hemibiotrophic pathogens. Increases in jasmonic acid (JA) synthesis near the sites of pathogen infection have also been described. [15] [16] This physiological response to increase JA production has been implicated in the ubiquitination of jasmonate ZIM domains (JAZ) proteins, which inhibit JA signaling, leading to their degradation and a subsequent increase in JA activated defense genes. [15]

Studies regarding the upregulation of defensive chemicals have confirmed the role of SA and JA in pathogen defense. In studies utilizing Arabidopsis mutants with the bacterial NahG gene, which inhibits the production and accumulation of SA, were shown to be more susceptible to pathogens than the wild-type plants. This was thought to result from the inability to produce critical defensive mechanisms including increased PR gene expression. [16] [17] Other studies conducted by injecting tobacco plants and Arabidopsis with salicylic acid resulted in higher resistance of infection by the alfalfa and tobacco mosaic viruses, indicating a role for SA biosynthesis in reducing viral replication. [17] [18] Additionally, studies performed using Arabidopsis with mutated jasmonic acid biosynthesis pathways have shown JA mutants to be at an increased risk of infection by soil pathogens. [16]

Along with SA and JA, other defensive chemicals have been implicated in plant viral pathogen defenses including abscisic acid (ABA), gibberellic acid (GA), auxin, and peptide hormones. [15] The use of hormones and innate immunity presents parallels between animal and plant defenses, though pattern-triggered immunity is thought to have arisen independently in each. [12]

Cross tolerance with abiotic stress

Remote sensing

The Agricultural Research Service (ARS) and various government agencies and private institutions have provided a great deal of fundamental information relating spectral reflectance and thermal emittance properties of soils and crops to their agronomic and biophysical characteristics. This knowledge has facilitated the development and use of various remote sensing methods for non-destructive monitoring of plant growth and development and for the detection of many environmental stresses that limit plant productivity. Coupled with rapid advances in computing and position locating technologies, remote sensing from ground-, air-, and space-based platforms is now capable of providing detailed spatial and temporal information on plant response to their local environment that is needed for site specific agricultural management approaches. [24] This is very important in today's society because with increasing pressure on global food productivity due to population increase, result in a demand for stress-tolerant crop varieties that has never been greater.

See also

Related Research Articles

<span class="mw-page-title-main">Calmodulin</span> Messenger protein

Calmodulin (CaM) (an abbreviation for calcium-modulated protein) is a multifunctional intermediate calcium-binding messenger protein expressed in all eukaryotic cells. It is an intracellular target of the secondary messenger Ca2+, and the binding of Ca2+ is required for the activation of calmodulin. Once bound to Ca2+, calmodulin acts as part of a calcium signal transduction pathway by modifying its interactions with various target proteins such as kinases or phosphatases.

<span class="mw-page-title-main">Plant hormone</span> Chemical compounds that regulate plant growth and development

Plant hormones are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, including embryogenesis, the regulation of organ size, pathogen defense, stress tolerance and reproductive development. Unlike in animals each plant cell is capable of producing hormones. Went and Thimann coined the term "phytohormone" and used it in the title of their 1937 book.

<span class="mw-page-title-main">Jasmonate</span> Lipid-based plant hormones

Jasmonate (JA) and its derivatives are lipid-based plant hormones that regulate a wide range of processes in plants, ranging from growth and photosynthesis to reproductive development. In particular, JAs are critical for plant defense against herbivory and plant responses to poor environmental conditions and other kinds of abiotic and biotic challenges. Some JAs can also be released as volatile organic compounds (VOCs) to permit communication between plants in anticipation of mutual dangers.

<span class="mw-page-title-main">Apoplast</span> Extracellular space, outside the cell membranes of plants

The apoplast is the extracellular space outside of plant cell membranes, especially the fluid-filled cell walls of adjacent cells where water and dissolved material can flow and diffuse freely. Fluid and material flows occurring in any extracellular space are called apoplastic flow or apoplastic transport. The apoplastic pathway is one route by which water and solutes are transported and distributed to different places through tissues and organs, contrasting with the symplastic pathway.

<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.

<span class="mw-page-title-main">Jasmonic acid</span> Chemical compound

Jasmonic acid (JA) is an organic compound found in several plants including jasmine. The molecule is a member of the jasmonate class of plant hormones. It is biosynthesized from linolenic acid by the octadecanoid pathway. It was first isolated in 1957 as the methyl ester of jasmonic acid by the Swiss chemist Édouard Demole and his colleagues.

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">Systemin</span> Plant peptide hormone

Systemin is a plant peptide hormone involved in the wound response in the family Solanaceae. It was the first plant hormone that was proven to be a peptide having been isolated from tomato leaves in 1991 by a group led by Clarence A. Ryan. Since then, other peptides with similar functions have been identified in tomato and outside of the Solanaceae. Hydroxyproline-rich glycopeptides were found in tobacco in 2001 and AtPeps were found in Arabidopsis thaliana in 2006. Their precursors are found both in the cytoplasm and cell walls of plant cells, upon insect damage, the precursors are processed to produce one or more mature peptides. The receptor for systemin was first thought to be the same as the brassinolide receptor but this is now uncertain. The signal transduction processes that occur after the peptides bind are similar to the cytokine-mediated inflammatory immune response in animals. Early experiments showed that systemin travelled around the plant after insects had damaged the plant, activating systemic acquired resistance, now it is thought that it increases the production of jasmonic acid causing the same result. The main function of systemins is to coordinate defensive responses against insect herbivores but they also affect plant development. Systemin induces the production of protease inhibitors which protect against insect herbivores, other peptides activate defensins and modify root growth. They have also been shown to affect plants' responses to salt stress and UV radiation. AtPEPs have been shown to affect resistance against oomycetes and may allow A. thaliana to distinguish between different pathogens. In Nicotiana attenuata, some of the peptides have stopped being involved in defensive roles and instead affect flower morphology.

<span class="mw-page-title-main">Leucyl aminopeptidase</span> Class of enzymes

Leucyl aminopeptidases are enzymes that preferentially catalyze the hydrolysis of leucine residues at the N-terminus of peptides and proteins. Other N-terminal residues can also be cleaved, however. LAPs have been found across superkingdoms. Identified LAPs include human LAP, bovine lens LAP, porcine LAP, Escherichia coli LAP, and the solanaceous-specific acidic LAP (LAP-A) in tomato.

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. 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).

<span class="mw-page-title-main">Plant disease resistance</span> Ability of plants to withstand pathogens

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.

In plant biology, elicitors are extrinsic or foreign molecules often associated with plant pests, diseases or synergistic organisms. Elicitor molecules can attach to special receptor proteins located on plant cell membranes. These receptors are able to recognize the molecular pattern of elicitors and trigger intracellular defence signalling via the octadecanoid pathway. This response results in the enhanced synthesis of metabolites which reduce damage and increase resistance to pest, disease or environmental stress. This is an immune response called pattern triggered immunity (PTI).

β-Aminobutyric acid Chemical compound

β-Aminobutyric acid (BABA) is an isomer of the amino acid aminobutyric acid with the chemical formula C4H9NO2. It has two isomers, α-aminobutyric acid and γ-aminobutyric acid (GABA), a neurotransmitter in animals that is also found in plants, where it may play a role in signalling. All three are non-proteinogenic amino acids, not being found in proteins. BABA is known for its ability to induce plant disease resistance, as well as increased resistance to abiotic stresses, when applied to plants.

WRKY transcription factors are proteins that bind DNA. They are transcription factors that regulate many processes in plants and algae (Viridiplantae), such as the responses to biotic and abiotic stresses, senescence, seed dormancy and seed germination and some developmental processes but also contribute to secondary metabolism.

Induced systemic resistance (ISR) is a resistance mechanism in plants that is activated by infection. Its mode of action does not depend on direct killing or inhibition of the invading pathogen, but rather on increasing physical or chemical barrier of the host plant. Like the Systemic Acquired Resistance (SAR) a plant can develop defenses against an invader such as a pathogen or parasite if an infection takes place. In contrast to SAR, which is triggered by the accumulation of salicylic acid, ISR instead relies on signal transduction pathways activated by jasmonate and ethylene.

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">Thimet oligopeptidase</span>

Thimet oligopeptidases, also known as TOPs, are a type of M3 metallopeptidases. These enzymes can be found in animals and plants, showing distinctive functions. In animals and humans, they are involved in the degradation of peptides, such as bradykinin, neurotensin, angiotensin I, and Aβ peptide, helping to regulate physiological processes. In plants, their role is related to the degradation of targeting peptides and the immune response to pathogens through Salicylic Acid (SA)-dependent stress signaling. In Arabidopsis thaliana—recognized as a model plant for scientific studies—two thimet oligopeptidases, known as TOP1 and TOP2, have been identified as targets for salicylic acid binding in the plant. These TOP enzymes are key components to understand the SA-mediated signaling where interactions exist with different components and most of the pathways are unknown.

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.

Plants are constantly exposed to different stresses that result in wounding. Plants have adapted to defend themselves against wounding events, like herbivore attacks or environmental stresses. There are many defense mechanisms that plants rely on to help fight off pathogens and subsequent infections. Wounding responses can be local, like the deposition of callose, and others are systemic, which involve a variety of hormones like jasmonic acid and abscisic acid.

Chemical defenses in <i>Cannabis</i> Defense of Cannabis plant from pathogens

Cannabis (/ˈkænəbɪs/) is commonly known as marijuana or hemp and has two known strains: Cannabis sativa and Cannabis indica, both of which produce chemicals to deter herbivory. The chemical composition includes specialized terpenes and cannabinoids, mainly tetrahydrocannabinol (THC), and cannabidiol (CBD). These substances play a role in defending the plant from pathogens including insects, fungi, viruses and bacteria. THC and CBD are stored mostly in the trichomes of the plant, and can cause psychological and physical impairment in the user, via the endocannabinoid system and unique receptors. THC increases dopamine levels in the brain, which attributes to the euphoric and relaxed feelings cannabis provides. As THC is a secondary metabolite, it poses no known effects towards plant development, growth, and reproduction. However, some studies show secondary metabolites such as cannabinoids, flavonoids, and terpenes are used as defense mechanisms against biotic and abiotic environmental stressors.

References

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  20. ↑ Perez & Brown 2014.
  21. ↑ Raju et al. 2015.
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