Isoprene synthesis in plants: lessons from a transgenic tobacco model.

Isoprene is a highly reactive gas, and is emitted in such large quantities from the biosphere that it substantially affects the oxidizing potential of the atmosphere. Relatively little is known about the control of isoprene emission at the molecular level. Using transgenic tobacco lines harbouring a poplar isoprene synthase gene, we examined control of isoprene emission. Isoprene synthase required chloroplastic localization for catalytic activity, and isoprene was produced via the methyl erythritol (MEP) pathway from recently assimilated carbon. Emission patterns in transgenic tobacco plants were remarkably similar to naturally emitting plants under a wide variety of conditions. Emissions correlated with photosynthetic rates in developing and mature leaves, and with the amount of isoprene synthase protein in mature leaves. Isoprene synthase protein levels did not change under short-term increase in heat/light, despite an increase in emissions under these conditions. A robust circadian pattern could be observed in emissions from long-day plants. The data support the idea that substrate supply and changes in enzyme kinetics (rather than changes in isoprene synthase levels or post-translational regulation of activity) are the primary controls on isoprene emission in mature transgenic tobacco leaves.

Biogenic volatile organic compounds in the Earth system.

Biogenic volatile organic compounds produced by plants are involved in plant growth, development, reproduction and defence. They also function as communication media within plant communities, between plants and between plants and insects. Because of the high chemical reactivity of many of these compounds, coupled with their large mass emission rates from vegetation into the atmosphere, they have significant effects on the chemical composition and physical characteristics of the atmosphere. Hence, biogenic volatile organic compounds mediate the relationship between the biosphere and the atmosphere. Alteration of this relationship by anthropogenically driven changes to the environment, including global climate change, may perturb these interactions and may lead to adverse and hard-to-predict consequences for the Earth system.

Isoprene synthesis protects transgenic tobacco plants from oxidative stress.

Isoprene emission represents a significant loss of carbon to those plant species that synthesize this highly volatile and reactive compound. As a tool for studying the role of isoprene in plant physiology and biochemistry, we developed transgenic tobacco plants capable of emitting isoprene in a similar manner to and at rates comparable to a naturally emitting species. Thermotolerance of photosynthesis against transient high-temperature episodes could only be observed in lines emitting high levels of isoprene; the effect was very mild and could only be identified over repetitive stress events. However, isoprene-emitting plants were highly resistant to ozone-induced oxidative damage compared with their non-emitting azygous controls. In ozone-treated plants, accumulation of toxic reactive oxygen species (ROS) was inhibited, and antioxidant levels were higher. Isoprene-emitting plants showed remarkably decreased foliar damage and higher rates of photosynthesis compared to non-emitting plants immediately following oxidative stress events. An inhibition of hydrogen peroxide accumulation in isoprene-emitting plants may stall the programmed cell death response which would otherwise lead to foliar necrosis. These results demonstrate that endogenously produced isoprene provides protection from oxidative damage.

Discrimination of plant volatile signatures by an electronic nose: aA potential technology for plant pest and disease monitoring.

The volatile organic compounds (VOCs) profile emitted from plants often changes in response to environmental factors, and monitoring the change of such profiles could provide a nondestructive means of plant health measurement An electronic nose (e-nose) was used to discriminate among VOC bouquets emitted by cucumber, pepper, and tomato leaves subjected to mechanical damage or pest and disease attacks compared with undamaged control leaves. Principle component analysis, discriminant function analysis, and cluster analysis were applied to evaluate the data. The results indicate that the e-nose can discriminate among VOCs from undamaged leaves of the three tested species. It can also discriminate undamaged and artificially damaged leaves of the same plant species. In cucumber, the e-nose can discriminate among VOCs emitted from control, artificially damaged, and spider-mite-infested leaves. It could also discriminate among VOCs emitted from control, artificially damaged, hornworm-damaged, and powdery-mildew-infected tomato leaves. The relationships between the changes in volatile signatures detected by the e-nose to changes in the underlying chemistry of plant VOC signatures in response to applied stresses were quantified by gas chromatography mass spectrometry. We conclude that the e-nose had genuine responses to changes in plant VOC signatures and can successfully discriminate them. These studies demonstrate the potential use of such e-nose technology as a real time pest and disease monitoring system in agricultural and horticultural settings.

Isoprene emissions influence herbivore feeding decisions.

Isoprene (C(5)H(8), 2-methyl 1,3-butadiene) is synthesized and emitted by many, but not all, plants. Unlike other related volatile organic compounds (monoterpenes and sesquiterpenes), isoprene has not been shown to mediate plant-herbivore interactions. Here, for the first time, we show, in feeding choice tests using isoprene-emitting transgenic tobacco plants (Nicotiana tabacum cv. Samsun) and non-emitting azygous control plants, that isoprene deters Manduca sexta caterpillars from feeding. This avoidance behaviour was confirmed using an artificial (isoprene-emitting and non-emitting control) diet. Both in vivo and in vitro experiments showed that isoprene can activate feeding avoidance behaviour in this system with a dose-response effect on caterpillar behaviour and an isoprene emission threshold level of <6 nmol m(-2) s(-1).

The role of isoprene in insect herbivory.

Several hypotheses have previously been proposed to explain the function of isoprene in plants, including its ability to protect the leaf metabolic machinery from transient high temperature1,2 and from oxidative stress.3 Isoprene may also serve as a metabolic overflow mechanism for carbon or photosynthetic energy4-6 and may promote flowering in neighbouring plants.7 We have reported recently that isoprene can be detected by a herbivore, Manduca sexta, and that it directly deters them from feeding, with an isoprene emission threshold level of <6 nmol m(-2) s(-1).8 We demonstrated this using both in vivo experiments, using isoprene-emitting transgenic tobacco plants (Nicotiana tabacum cv. Samsun) and non-emitting azygous control plants, and in vitro experiments, using an artificial (isoprene-emitting and non-emitting control) diet. Here we discuss the potential role of isoprene in plant-herbivore interactions and the possibility that isoprene actually serves multiple purposes in plants.