Green leaf volatiles (GLV) are organic compounds released by plants. [1] Some of these chemicals function as signaling compounds between either plants of the same species, of other species, or even different lifeforms like insects. [2] [3] [4] [5]
Green leaf volatiles are involved in patterns of attack and protection between species. They have been found to increase the attractive effect of pheromones of cohabiting insect species that protect plants from attacking insect species. For example, corn plants that are being fed on by caterpillars will release GLVs that attract wasps, who then attack the caterpillars. [2] [4] GLVs also have antimicrobial properties that can prevent infection at the site of injury. [3]
GLVs include C6-aldehydes [(Z)-3-hexenal, n-hexanal] and their derivatives such as (Z)-3-hexenol, (Z)-3-hexen-1-yl acetate, and the corresponding E-isomers. [6] [7]
When a plant is attacked, it emits GLVs into the environment through the air. How a plant responds depends on the type of damage involved. Plants respond differently to damage from a purely mechanical source and damage from herbivores. Mechanical damage tends to cause damage-associated molecular patterns (DAMPs) involving plant-derived substances and breakdown products. Herbivore-associated molecular patterns (HAMPs) involve characteristic molecules left by different types of herbivores when feeding. The oral secretions of herbivores appear to play an essential role in triggering the release of species-specific herbivore-induced plant volatiles. Wounds from herbivores, and mechanical wounds that have been treated with herbivore oral secretions, both trigger the release of higher quantities of plant volatiles than mechanical damage. [4]
Volatile blends are proposed to convey a variety of information to insects and plants. "Each plant species and even each plant genotype releases its own specific blend, and the quantities and ratios in which they are released also vary with the arthropod that is feeding on a plant and may even provide information on the time of day that feeding occurs." [4] In addition to GLVs, herbivore induced plant volatiles (HIPVs) include terpenes, ethylene, methyl salicylate and other VOCs. [6]
GLVs activate the expression of genes related to the plants' defense mechanisms. [3] Different antagonists trigger different expression of genes and the biosynthesis of signaling peptides which mediate systemic defense responses. [4]
Undamaged neighboring plants have been shown, in some cases, to respond to GLV signals. [3] Both the plant emitting the GLVs and its neighboring plants can enter a primed state in which plants activate their defenses systems more quickly and in a stronger concentration. [8] [4]
The first study to clearly demonstrate anti-herbivore defense priming by GLVs focused on corn ( Zea mays ). Neighboring plants responded to the release of GLVs by priming against insect herbivore attack, reacting more rapidly and releasing greater levels of GLVs. [3] [9] Similar results have been shown in tomato plants. Neighboring plants reacted more strongly to GLVs from the plants exposed to the herbivore, by releasing more of the proteins related to the plants' defense mechanisms. [10]
In positive plant-insect interactions, GLVs are used as a form of defense. They attract predators to plants that are being preyed upon by herbivores. [4] For example, female parasitoid wasps from two different families, Microplitis croceipes and Netelia heroica , can be attracted to plants that are emitting GLVs due to wounding from caterpillars. [11] Maize plants emit volatiles to attract the parasitic wasps Cotesia marginiventris and Microplitis rufiventris to attack African cotton leafworm. [12] [13] In some species GLVs enhance the attraction of sex pheromones. [4] [14] For example, green leaf volatiles have been found to increase the response of tobacco budworm to sex pheromone. Budworm larvae feed on tobacco, cotton, and various flowers and weeds, and in turn can be fed on by the larvae of cohabiting species that are attracted by GLVs. [15]
In another study, a multi-plant relationship was reported. The parasitic wasps ( Vespula germanica and V. vulgaris ) prey on caterpillar (Pieris brassicae)-infested cabbage leaves that emit GLVs. The same GLVs are emitted by the orchids ( Epipactis purpurata and E. helleborine ) through pheromone release. The orchids benefit from attracting the wasps, not to protect them from insects, but because the wasps aid in pollination. [16] [17]
Benefits of GLV release have also been reported in soybeans grown in Iowa. [18] When these soybean plants became heavily infested by aphids, the amount of GLV released far surpassed normal levels and as a result, more spotted lady beetles were attracted to the pheromone releasing plants and preyed on the bugs eating the plant. The stimulus of aphid predation is chemically transmitted through the plant to coordinate an increase release of GLV’s. The particular chemical released is unique to these spotted lady beetles and when different species of beetles were tested, there wasn’t any extra inclination for them to move towards GLV releasing plants. [18] This indicates that these soybeans evolved ability to release species-specific pheromones to aid in their survival.
GLV release is correlated with fruit ripeness. [19] Although this may be of effect in attracting pollinators, it also can cause issues if these GLV’s attract predators. One such example of this is with boll weevils, as an increase of GLV release when the plants are ripe has been found to increase the predation rate of these beetles. [19]
Another issue with GLV release and increasing predation is with populations that alter GLV emissions from the affected plants. In one case, it was noted that secretions from certain species of caterpillars significantly decrease the effect amount of GLV emissions. [20] In order to determine what was being done to decrease GLV emissions, a study was run on four unique species of caterpillars to measure their effectiveness in decreasing GLV levels released from the predated plant. It’s been found that compounds in the gut and salivary glands, as well as modifications to those compounds in these various species, has been able to mute a large part of the effect of GLV released into the external environment. [20] How this is done is though stopping the flow of pheromone molecules, so they can’t interact with receptors on the leaves of other plants. [20]
GLVs can also have antimicrobial effects. [21] Some plants express HPL, the main enzyme of GLV synthesis. [8] The rates of fungal spore growth in HPL over-expressing have been compared with HPL silencing mutants to the wild type plants. [8] Results from the study showed lower rates of fungal growth and higher GLV emissions on the HPL over-expressing mutants, while the HPL silencing mutants showed higher rates of fungal growth and lower GLV emissions, which supports the hypothesis that GLVs have antimicrobial properties. [8]
The antimicrobial properties of GLVs have also been proposed to be part of an evolutionary arms race. During an infection, plants emit GLVs to act as microbial agents, but bacteria and viruses have adapted to use these GLVs to their own benefit. [22] The most common example of this is found in the red raspberry. When the red raspberry plant is infected, the virus influences it to produce more GLVs, which attract the red raspberry aphid. [9] These GLVs cause more aphids to come and to feed on the plant for longer, giving the virus better chances of being ingested and spread more widely. [9] Researchers are now trying to determine whether under infectious conditions plants emit GLVs for their benefit, or if bacteria and viruses induce the release of these compounds for their own benefit. [23] Studies in this area have been inconclusive and contradictory.
A systematic review by Schuman 2023 finds that most studies on plant volatiles relate to herbivore interactions. Schuman also finds that laboratory studies are overrepresented despite the wide differences in herbivore behaviour between natural and artificial settings. [25]
Companion planting in gardening and agriculture is the planting of different crops in proximity for any of a number of different reasons, including weed suppression, pest control, pollination, providing habitat for beneficial insects, maximizing use of space, and to otherwise increase crop productivity. Companion planting is a form of polyculture.
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Chemical ecology is a vast and interdisciplinary field utilizing biochemistry, biology, ecology, and organic chemistry for explaining observed interactions of living things and their environment through chemical compounds. Early examples of the field trace back to experiments with the same plant genus in different environments, interaction of plants and butterflies, and the behavioral effect of catnip. Chemical ecologists seek to identify the specific molecules that function as signals mediating community or ecosystem processes and to understand the evolution of these signals. The chemicals behind such roles are typically small, readily-diffusible organic molecules that act over various distances that are dependent on the environment but can also include larger molecules and small peptides.
A hyperparasite, also known as a metaparasite, is a parasite whose host, often an insect, is also a parasite, often specifically a parasitoid. Hyperparasites are found mainly among the wasp-waisted Apocrita within the Hymenoptera, and in two other insect orders, the Diptera and Coleoptera (beetles). Seventeen families in Hymenoptera and a few species of Diptera and Coleoptera are hyperparasitic. Hyperparasitism developed from primary parasitism, which evolved in the Jurassic period in the Hymenoptera. Hyperparasitism intrigues entomologists because of its multidisciplinary relationship to evolution, ecology, behavior, biological control, taxonomy, and mathematical models.
A semiochemical, from the Greek σημεῖον (semeion), meaning "signal", is a chemical substance or mixture released by an organism that affects the behaviors of other individuals. Semiochemical communication can be divided into two broad classes: communication between individuals of the same species (intraspecific) or communication between different species (interspecific).
A kairomone is a semiochemical released by an organism that mediates interspecific interactions in a way that benefits a different species at the expense of the emitter. Derived from the Greek καιρός, meaning "opportune moment", it serves as a form of "eavesdropping", enabling the receiver to gain an advantage, such as locating food or evading predators, even if it poses a risk to the emitter. Unlike allomones, which benefit the producer at the receiver's cost, or synomones, which are mutually beneficial, kairomones favor only the recipient. Primarily studied in entomology, kairomones can play key roles in predator-prey dynamics, mate attraction, and even applications in pest control.
Nectar is a viscous, sugar-rich liquid produced by plants in glands called nectaries, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, which in turn provide herbivore protection. Common nectar-consuming pollinators include mosquitoes, hoverflies, wasps, bees, butterflies and moths, hummingbirds, honeyeaters and bats. Nectar is an economically important substance as it is the sugar source for honey. It is also useful in agriculture and horticulture because the adult stages of some predatory insects feed on nectar. For example, a number of predacious or parasitoid wasps rely on nectar as a primary food source. In turn, these wasps then hunt agricultural pest insects as food for their young.
Plant defense against herbivory or host-plant resistance is a range of adaptations evolved by plants which improve their survival and reproduction by reducing the impact of herbivores. Many plants produce secondary metabolites, known as allelochemicals, that influence the behavior, growth, or survival of herbivores. These chemical defenses can act as repellents or toxins to herbivores or reduce plant digestibility. Another defensive strategy of plants is changing their attractiveness. Plants can sense being touched, and they can respond with strategies to defend against herbivores. To prevent overconsumption by large herbivores, plants alter their appearance by changing their size or quality, reducing the rate at which they are consumed.
Insect ecology is the interaction of insects, individually or as a community, with the surrounding environment or ecosystem. This interaction is mostly mediated by the secretion and detection of chemicals (semiochemical) in the environment by insects. Semiochemicals are secreted by the organisms in the environment and they are detected by other organism such as insects. Semiochemicals used by organisms, including (insects) to interact with other organism either of the same species or different species can generally grouped into four. These are pheromone, synomones, allomone and kairomone. Pheromones are semiochemicals that facilitates interaction between organisms of same species. Synomones benefit both the producer and receiver, allomone is advantageous to only the producer whiles kairomones is beneficial to the receiver. Insect interact with other species within their community and these interaction include mutualism, commensalism, ammensalism, parasitism and neutralisms.
Nicotiana attenuata is a species of wild tobacco known by the common name coyote tobacco. It is native to western North America from British Columbia to Texas and northern Mexico, where it grows in many types of habitat. It is a glandular and sparsely hairy annual herb exceeding a meter in maximum height. The leaf blades may be 10 centimetres (4 in) long, the lower ones oval and the upper narrower in shape, and are borne on petioles. The inflorescence bears several flowers with pinkish or greenish white tubular throats 2 to 3 centimetres long, their bases enclosed in pointed sepals. The flower face has five mostly white lobes. The fruit is a capsule about 1 centimetre long.
Chemical defense is a strategy employed by many organisms to avoid consumption by producing toxic or repellent metabolites or chemical warnings which incite defensive behavioral changes. The production of defensive chemicals occurs in plants, fungi, and bacteria, as well as invertebrate and vertebrate animals. The class of chemicals produced by organisms that are considered defensive may be considered in a strict sense to only apply to those aiding an organism in escaping herbivory or predation. However, the distinction between types of chemical interaction is subjective and defensive chemicals may also be considered to protect against reduced fitness by pests, parasites, and competitors. Repellent rather than toxic metabolites are allomones, a sub category signaling metabolites known as semiochemicals. Many chemicals used for defensive purposes are secondary metabolites derived from primary metabolites which serve a physiological purpose in the organism. Secondary metabolites produced by plants are consumed and sequestered by a variety of arthropods and, in turn, toxins found in some amphibians, snakes, and even birds can be traced back to arthropod prey. There are a variety of special cases for considering mammalian antipredatory adaptations as chemical defenses as well.
Insects have a wide variety of predators, including birds, reptiles, amphibians, mammals, carnivorous plants, and other arthropods. The great majority (80–99.99%) of individuals born do not survive to reproductive age, with perhaps 50% of this mortality rate attributed to predation. In order to deal with this ongoing escapist battle, insects have evolved a wide range of defense mechanisms. The only restraint on these adaptations is that their cost, in terms of time and energy, does not exceed the benefit that they provide to the organism. The further that a feature tips the balance towards beneficial, the more likely that selection will act upon the trait, passing it down to further generations. The opposite also holds true; defenses that are too costly will have a little chance of being passed down. Examples of defenses that have withstood the test of time include hiding, escape by flight or running, and firmly holding ground to fight as well as producing chemicals and social structures that help prevent predation.
Rhopalosiphum maidis, common names corn leaf aphid and corn aphid, is an insect, and a pest of maize and other crops. It has a nearly worldwide distribution and is typically found in agricultural fields, grasslands, and forest-grassland zones. Among aphids that feed on maize, it is the most commonly encountered and most economically damaging, particularly in tropical and warmer temperate areas. In addition to maize, R. maidis damages rice, sorghum, and other cultivated and wild monocots.
In evolutionary biology, mimicry in plants is where a plant evolves to resemble another organism physically or chemically. Mimicry in plants has been studied far less than mimicry in animals. It may provide protection against herbivory, or may deceptively encourage mutualists, like pollinators, to provide a service without offering a reward in return.
Tritrophic interactions in plant defense against herbivory describe the ecological impacts of three trophic levels on each other: the plant, the herbivore, and its natural enemies. They may also be called multitrophic interactions when further trophic levels, such as soil microbes, endophytes, or hyperparasitoids are considered. Tritrophic interactions join pollination and seed dispersal as vital biological functions which plants perform via cooperation with animals.
Floral scent, or flower scent, is composed of all the volatile organic compounds (VOCs), or aroma compounds, emitted by floral tissue. Other names for floral scent include, aroma, fragrance, floral odour or perfume. Flower scent of most flowering plant species encompasses a diversity of VOCs, sometimes up to several hundred different compounds. The primary functions of floral scent are to deter herbivores and especially folivorous insects, and to attract pollinators. Floral scent is one of the most important communication channels mediating plant-pollinator interactions, along with visual cues.
A phytobiome consists of a plant (phyto) situated in its specific ecological area (biome), including its environment and the associated communities of organisms which inhabit it. These organisms include all macro- and micro-organisms living in, on, or around the plant including bacteria, archaea, fungi, protists, insects, animals, and other plants. The environment includes the soil, air, and climate. Examples of ecological areas are fields, rangelands, forests. Knowledge of the interactions within a phytobiome can be used to create tools for agriculture, crop management, increased health, preservation, productivity, and sustainability of cropping and forest systems.
Plants are exposed to many stress factors such as disease, temperature changes, herbivory, injury and more. Therefore, in order to respond or be ready for any kind of physiological state, they need to develop some sort of system for their survival in the moment and/or for the future. Plant communication encompasses communication using volatile organic compounds, electrical signaling, and common mycorrhizal networks between plants and a host of other organisms such as soil microbes, other plants, animals, insects, and fungi. Plants communicate through a host of volatile organic compounds (VOCs) that can be separated into four broad categories, each the product of distinct chemical pathways: fatty acid derivatives, phenylpropanoids/benzenoids, amino acid derivatives, and terpenoids. Due to the physical/chemical constraints most VOCs are of low molecular mass, are hydrophobic, and have high vapor pressures. The responses of organisms to plant emitted VOCs varies from attracting the predator of a specific herbivore to reduce mechanical damage inflicted on the plant to the induction of chemical defenses of a neighboring plant before it is being attacked. In addition, the host of VOCs emitted varies from plant to plant, where for example, the Venus Fly Trap can emit VOCs to specifically target and attract starved prey. While these VOCs typically lead to increased resistance to herbivory in neighboring plants, there is no clear benefit to the emitting plant in helping nearby plants. As such, whether neighboring plants have evolved the capability to "eavesdrop" or whether there is an unknown tradeoff occurring is subject to much scientific debate. As related to the aspect of meaning-making, the field is also identified as phytosemiotics.
Bergamotenes are a group of isomeric chemical compounds with the molecular formula C15H24. The bergamotenes are found in a variety of plants, particularly in their essential oils.
The smell of freshly cut grass is an odour caused by green leaf volatiles (GLVs) released when it is damaged. Mechanical damage to grass from activities such as lawnmowing results in the release of cis-3-hexenal and other compounds that contribute to a grassy or "green" smell. cis-3-Hexenal has a low odour detection threshold that humans can perceive at concentrations as low as 0.25 parts per billion.