Plant communication

The terpenoid verbenone is a plant pheromone, signalling to insects that a tree is already infested by beetles.[13]

Terpenoids facilitate communication between plants and insects, mammals, fungi, microorganisms, and other plants.[14] Terpenoids may act as both attractants and repellants for various insects. For example, pine shoot beetles (Tomicus piniperda) are attracted to certain monoterpenes ( (+/-)-a-pinene, (+)-3-carene and terpinolene) produced by Scots pines (Pinus sylvestris), while being repelled by others (such as verbenone).[15]

Terpenoids are a large family of biological molecules with over 22,000 compounds.[16] Terpenoids are similar to terpenes in their carbon skeleton but unlike terpenes contain functional groups. The structure of terpenoids is described by the biogenetic isoprene rule which states that terpenoids can be thought of being made of isoprenoid subunits, arranged either regularly or irregularly.[17] The biosynthesis of terpenoids occurs via the methylerythritol phosphate (MEP) and mevalonic acid(MVA) pathways[6] both of which include isopentenyl diphosphate (IPP) and dimethylallyl diphosphate (DMAPP) as key components.[18] The MEP pathway produces hemiterpenes, monoterpenes, diterpenes, and volatile carotenoid derivatives while the MVA pathway produces sesquiterpenes.[6]

Pisum sativum (garden pea) plants communicate stress cues via their roots to allow neighboring unstressed plants to anticipate an abiotic stressor. Pea plants are commonly grown in temperate regions throughout the world.[19] However, this adaptation allows plants to anticipate abiotic stresses such as drought. In 2011, Falik et al. tested the ability of unstressed pea plants to sense and respond to stress cues by inducing osmotic stress on a neighboring plant.[20] Falik et al. subjected the root of an externally-induced plant to mannitol in order to inflict osmotic stress and drought-like conditions. Five unstressed plants neighbored both sides of this stressed plant. On one side, the unstressed plants shared their root system with their neighbors to allow for root communication. On the other side, the unstressed plants did not share root systems with their neighbors.[20]

Falik et al. found that unstressed plants demonstrated the ability to sense and respond to stress cues emitted from the roots of the osmotically stressed plant. Furthermore, the unstressed plants were able to send additional stress cues to other neighboring unstressed plants in order to relay the signal. A cascade effect of stomatal closure was observed in neighboring unstressed plants that shared their rooting system but was not observed in the unstressed plants that did not share their rooting system.[20] Therefore, neighboring plants demonstrate the ability to sense, integrate, and respond to stress cues transmitted through roots. Although Falik et al. did not identify the chemical responsible for perceiving stress cues, research conducted in 2016 by Delory et al. suggests several possibilities. They found that plant roots synthesize and release a wide array of organic compounds including solutes and volatiles (i.e. terpenes).[21] They cited additional research demonstrating that root-emitted molecules have the potential to induce physiological responses in neighboring plants either directly or indirectly by modifying the soil chemistry.[21] Moreover, Kegge et al. demonstrated that plants perceive the presence of neighbors through changes in water/nutrient availability, root exudates, and soil microorganisms.[22]

Although the underlying mechanism behind stress cues emitted by roots remains largely unknown, Falik et al. suggested that the plant hormone abscisic acid (ABA) may be responsible for integrating the observed phenotypic response (stomatal closure).[20] Further research is needed to identify a well-defined mechanism and the potential adaptive implications for priming neighbors in preparation for forthcoming abiotic stresses; however, a literature review by Robbins et al. published in 2014 characterized the root endodermis as a signaling control center in response to abiotic environmental stresses including drought.[23] They found that the plant hormone ABA regulates the root endodermal response under certain environmental conditions. In 2016 Rowe et al. experimentally validated this claim by showing that ABA regulated root growth under osmotic stress conditions.[24] Additionally, changes in cytosolic calcium concentrations act as signals to close stomata in response to drought stress cues. Therefore, the flux of solutes, volatiles, hormones, and ions are likely involved in the integration of the response to stress cues emitted by roots.

Another form of plant communication occurs through their complex root networks. Through roots, plants can share many different resources including nitrogen, fungi, nutrients, microbes, and carbon. This transfer of below ground carbon is examined in Philip et al. 2011. The goals of this paper were to test if carbon transfer was bi-directional, if one species had a net gain in carbon, and if more carbon was transferred through the soil pathway or common mycorrhizal network (CMN). CMNs occur when fungal mycelia link roots of plants together.[25] To test this, the researchers followed seedlings of paper birch and Douglas-fir in a greenhouse for 8 months, where hyphal linkages that crossed their roots were either severed or left intact. The experiment measured amounts of CO2 in both seedlings. It was discovered that there was indeed a bi-directional sharing of CO2 between the two trees, with the Douglas-fir receiving a slight net gain in CO2. Also, the carbon was transferred through both soil and the CMN pathways, as transfer occurred when the CMN linkages were interrupted, but much more transfer occurred when the CMN's were left unbroken. This experiment showed that through fungal mycelia linkage of the roots of two plants, plants are able to communicate with one another and transfer nutrients as well as other resources through below ground root networks.[25] Further studies go on to argue that this underground “tree talk” is crucial in the adaptation of forest ecosystems. Plant genotypes have shown that mycorrhizal fungal traits are heritable and play a role in plant behavior. These relationships with fungal networks can be mutualistic, commensal, or even parasitic. It has been shown that plants can rapidly change behavior such as root growth, shoot growth, photosynthetic rate, and defense mechanisms in response to mycorrhizal colonization.[26] Through root systems and common mycorrhizal networks, plants are able to communicate with one another below ground and alter behaviors or even share nutrients depending on different environmental cues.