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Some insectivorous birds orient towards insect‐defoliated trees even when they do not see the foliar damage or the herbivores. There are, however, only a few studies that have examined the mechanisms behind this foraging behaviour. Previous studies suggest that birds can use olfactory foraging cues (e.g. volatile organic compounds (VOCs) emitted by defoliated plants), indirect visual cues or a combination of the two sensory cues. VOCs from insect‐defoliated plants are known to attract natural enemies of herbivores, and researchers have hypothesized that VOCs could also act as olfactory foraging cues for birds. We conducted three experiments across a range of spatial scales to test this hypothesis. In each experiment, birds were presented with olfactory cues and their behavioural responses or foraging outcomes were observed. In the first experiment, two different VOC blends, designed to simulate the volatile emissions of mountain birch (Betula pubescens ssp. czerepanovii) after defoliation by autumnal moth (Epirrita autumnata) larvae, were used in behavioural experiments in aviaries with pied flycatchers (Ficedula hypoleuca). The second experiment was a field‐based trial of bird foraging efficiency; the same VOC blends were applied to mountain birches, silver birches (B. pendula) and European white birches (B. pubescens) with plasticine larvae attached to the trees to serve as artificial prey for birds and provide a means to monitor predation rate. In the third experiment, the attractiveness of silver birch saplings defoliated by autumnal moth larvae versus intact controls was tested with great tits (Parus major) and blue tits (Cyanistes caeruleus) in an aviary. Birds did not orient towards either artificial or real trees with VOC supplements or towards herbivore‐damaged saplings when these saplings and undamaged alternatives were hidden from view. These findings do not support the hypothesis that olfactory foraging cues are necessary in the attraction of birds to herbivore‐damaged trees.

One of the first observations that plants might utilize cues released by damaged neighbors under natural conditions was made in birch forests in 1985. However, the mechanisms underlying the observations were not determined, and birch (Betula spp.) has been neglected as a study system for inter-plant interaction ever since. Volatiles released by vegetative plant parts in response to herbivore damage play important roles as signals in plant-to-plant interactions in a range of woody and herbaceous plant species, and also have been shown to mediate signaling between branches of the same plant that have limited vascular connection. We established greenhouse experiments to assess: 1) whether exposure to plant volatiles from herbivore-damaged birches primes defense responses in undamaged neighbors; and 2) whether defenses also are primed in undamaged parts of the same plants with limited vascular connection. We observed a priming of defense responses, which were manifested in an augmented emission of terpenes and aromatic compounds in undamaged conspecific neighbors, and also an augmented emission of green leaf volatiles in systemic branches. Our work provides strong evidence of inter-plant signaling by volatiles, and an intra-plant systemic response in birch. However, the responses are specific, with emissions of different groups of plant volatiles typifying the primed response. This work complements and extends the previous work conducted with a natural population of birches.

We examined the effects of defoliation concurrently with elevated temperature and CO 2 on some chemical and morphological characteristics in the leaves of silver birch seedlings (Betula pendula). We also analyzed the consequent changes in the palatability of leaves for adult blue alder leaf beetles (Agelastica alni). Under the different climatic treatments, the seedlings were subjected to three fertilizer treatments (0 kg, 130 kg and 270 kg N ha -1 ) and defoliation treatments (0 %, 25 % and 50 % of the total leaf area). In each climatic treatment, fertilization increased the nitrogen content in the leaves, but decreased total concentrations of soluble phenolics, detected by high-performance liquid chromatography (HPLC), and insoluble condensed tannins. Defoliation, both independently and in combination with elevated temperature and CO 2 , decreased the concentrations of the phenolics. Compared to the intact controls, the leaves of the defoliated seedlings were smaller and tougher. Under elevated temperature, the beetles consumed a smaller amount of the leaves of plants subjected to the high fertilization, while under ambient climatic conditions, fertilization increased the feeding. The total leaf consumption was higher under the ambient climatic conditions than under elevated temperature, elevated CO 2 or the combination of elevated temperature and CO 2 .

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