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28 Communication Between Undamaged Plants by Volatiles 427<br />

In laboratory experiments, barley responses to allelobiotic interactions<br />

via volatiles from a different barley cultivar were measured in terms of<br />

changes in leaf temperature using an infrared camera (Pettersson et al.<br />

1999). All possible pairwise combinations of four barley cultivars were<br />

tested. When certain cultivars were exposed to volatiles from certain other<br />

cultivars, significant reductions in leaf temperature were found.<br />

Changes in leaf temperature result from active regulation of stomatal<br />

aperture transpiration, which depends on the water status of the plant<br />

modified by prevailing environmental conditions. Stomatal conductance,<br />

in turn, is a measure of transpiration. It has been reported that infrared<br />

measurements correlate well with data obtained using a diffusion porometer<br />

and with gas-exchange measurements, and studies have also shown local<br />

leaf temperature increases in regions where stomatal closure was induced<br />

by initial virus infection, before disease symptoms were visible (Chaerle<br />

and van der Streaten 2000).<br />

Reduction in barley leaf temperature following allelobiosis indicates<br />

increased transpiration rates in exposed plants. It is probable that an<br />

increased need for water necessitates increased allocation of available<br />

biomass to roots in these plants. Higher stomatal conductance enhances<br />

the influx of CO2, which is required to maintain a higher photosynthetic<br />

activity. The ULR is a physiological component of the RGR that is generally<br />

strongly correlated with the rate of photosynthesis (Poorter and Nagel<br />

2000). To maintain the same level of the ULR, it seems that the photosynthetic<br />

activity of allelobiosis-exposed plants increased.<br />

28.3<br />

Allelobiosis and Insect Responses<br />

Studies on the effects of plant–plant communication on insects have focused<br />

almost exclusively on interactions in which the responding plant is<br />

exposed to volatiles from herbivore- or pathogen-attacked plants. Volatiles<br />

produced by plants attacked in this way can induce responses in neighbouring<br />

undamaged plants, making them less attractive to herbivores (Bruin<br />

and Dicke 2001) and more attractive to the herbivores’ natural enemies<br />

(Dicke and Van Loon 2000). Recent studies at the biochemical and genetic<br />

level have started to clarify the set of changes induced in responding<br />

plants by exposure to volatiles from herbivore- or pathogen-attacked plants<br />

(Arimura et al. 2000; Farmer 2001; Pickett et al. 2001). However, volatile<br />

communication between undamaged plants, and its implications for higher<br />

trophic levels, i.e. insect herbivores and their natural enemies, has been less<br />

studied.

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