Who is my neighbor? Volatile cues in plant interactions.

One of the most important challenges for individual plants is coexistence with their neighbors. To compensate for their sessile lifestyle, plants developed complex and sophisticated chemical systems of communication among each other. Site-specific biotic and abiotic factors constantly alter the physiological activity of plants, which causes them to release various secondary metabolites in their environments. Volatile organic compounds (VOCs) are the most common cues that reflect a plant's current physiological status. In this sense, the identity of its immediate neighbors may have the greatest impact for a plant, as they share the same available resources. Plants constantly monitor and respond to these cues with great sensitivity and discrimination, resulting in specific changes in their growth pattern and adjusting their physiology, morphology, and phenotype accordingly. Those typical competition responses in receivers may increase their fitness as they can be elicited even before the competition takes place. Plant-plant interactions are dynamic and complex as they can include many different and important surrounding cues. A major challenge for all individual plants is detecting and actively responding only to "true" cues that point to real upcoming threat. Such selective responses to highly specific cues embedded in volatile bouquets are of great ecological importance in understanding plant-plant interactions. We have reviewed recent research on the role of VOCs in complex plant-plant interactions in plant-cross kingdom and highlighted their influence on organisms at higher trophic levels.

Pest suppression in cultivar mixtures is influenced by neighbor-specific plant-plant communication.

Increased plant genotypic diversity in crop fields can promote ecosystem services including pest control, but understanding of mechanisms behind herbivore population responses to cultivar mixtures is limited. We studied aphid settling on barley plants exposed to volatiles from different cultivars, aphid population development in monocultures and two-cultivar mixtures, and differences in volatile composition between studied cultivars. Aphid responses to one cultivar in a mixture were neighbor-specific and this was more important for pest suppression than the overall mixture effect, aphid colonization patterns, or natural enemy abundance. Aphid populations decreased most in a mixture where both cultivars showed a reduced aphid-plant acceptance after reciprocal volatile exposure in the laboratory, and reduced population growth compared to monocultures in the field. Our findings suggest that herbivore population responses to crop genotypic diversity can depend on plant-plant volatile interactions, which can lead to changes in herbivore response to individual cultivars in a mixture, resulting in slower population growth. The impact of plant-plant interaction through volatiles on associated herbivore species is rarely considered, but improved understanding of these mechanisms would advance our understanding of the ecological consequences of biodiversity and guide development of sustainable agricultural practices. Combining cultivars in mixtures based on how they interact with each other is a promising strategy for sustainable pest management.

Volatile interaction between undamaged plants affects tritrophic interactions through changed plant volatile emission.

Volatile interactions between unattacked plants can lead to changes in their volatile emissions. Exposure of potato plants to onion plant volatiles results in increased emission of 2 terpenoids, (E)-nerolidol and TMTT. We investigated whether this is detectable by the ladybird Coccinella septempunctata. The odor of onion-exposed potato was significantly more attractive to ladybirds than that of unexposed potato. Further, a synthetic blend mimicking the volatile profile of onion-exposed potato was more attractive than a blend mimicking that of unexposed potato. When presented individually, TMTT was attractive to ladybirds whereas (E)-nerolidol was repellent. Volatile exchange between unattacked plants and consequent increased attractiveness for ladybirds may be a mechanism that contributes to the increased abundance of natural enemies in complex plant habitats.

Volatile exchange between undamaged plants - a new mechanism affecting insect orientation in intercropping.

Changes in plant volatile emission can be induced by exposure to volatiles from neighbouring insect-attacked plants. However, plants are also exposed to volatiles from unattacked neighbours, and the consequences of this have not been explored. We investigated whether volatile exchange between undamaged plants affects volatile emission and plant-insect interaction. Consistently greater quantities of two terpenoids were found in the headspace of potato previously exposed to volatiles from undamaged onion plants identified by mass spectrometry. Using live plants and synthetic blends mimicking exposed and unexposed potato, we tested the olfactory response of winged aphids, Myzus persicae. The altered potato volatile profile deterred aphids in laboratory experiments. Further, we show that growing potato together with onion in the field reduces the abundance of winged, host-seeking aphids. Our study broadens the ecological significance of the phenomenon; volatiles carry not only information on whether or not neighbouring plants are under attack, but also information on the emitter plants themselves. In this way responding plants could obtain information on whether the neighbouring plant is a competitive threat and can accordingly adjust their growth towards it. We interpret this as a response in the process of adaptation towards neighbouring plants. Furthermore, these physiological changes in the responding plants have significant ecological impact, as behaviour of aphids was affected. Since herbivore host plants are potentially under constant exposure to these volatiles, our study has major implications for the understanding of how mechanisms within plant communities affect insects. This knowledge could be used to improve plant protection and increase scientific understanding of communication between plants and its impact on other organisms.