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Adaptive plant responses to specific abiotic stresses or biotic agents are fine-tuned by a network of hormonal signaling cascades, including abscisic acid (ABA), ethylene, jasmonic acid (JA) and salicylic acid. Moreover, hormonal cross-talk modulates plant responses to abiotic stresses and defenses against insect herbivores when they occur simultaneously. How such interactions affect plant responses under multiple stresses, however, is less understood, even though this may frequently occur in natural environments. Here, we review our current knowledge on how hormonal signaling regulates abiotic stress responses and defenses against insects, and discuss the few recent studies that attempted to dissect hormonal interactions occurring under simultaneous abiotic stress and herbivory. Based on this we hypothesize that drought stress enhances insect resistance due to synergistic interactions between JA and ABA signaling. Responses to flooding or waterlogging involve ethylene signaling, which likely reduces plant resistance to chewing herbivores due to its negative cross-talk with JA. However, the outcome of interactions between biotic and abiotic stress signaling is often plant and/or insect species-dependent and cannot simply be predicted based on general knowledge on the involvement of signaling pathways in single stress responses. More experimental data on non-model plant and insect species are needed to reveal general patterns and better understand the molecular mechanisms allowing plants to optimize their responses in complex environments.

Key message Pollen heat acclimation. Abstract As a consequence of global warming, plants have to face more severe and more frequently occurring periods of high temperature stress. While this affects the whole plant, development of the male gametophyte, the pollen, seems to be the most sensitive process. Given the great importance of functioning pollen for the plant life cycle and for agricultural production, it is necessary to understand this sensitivity. While changes in temperature affect different components of all cells and require a cellular response and acclimation, high temperature effects and responses in developing pollen are distinct from vegetative tissues at several points. This could be related to specific physiological characteristics of developing pollen and supporting tissues which make them vulnerable to high temperature, or its derived effects such as ROS accumulation and carbohydrate starvation. But also expression of heat stress-responsive genes shows unique patterns in developing pollen when compared to vegetative tissues that might explain the failure to withstand high temperatures. As an alternative to viewing pollen failure under high temperature as a result of inherent sensitivity of a specific developmental process, we end by discussing whether it might actually be an adaptation.

High ambient temperature has adverse effects on plant vegetative and reproductive development and reduces crop yield. To better understand the importance of male and female fertility for tomato fruit set ability under high temperature conditions and to test whether heat tolerance levels among and between reproductive and vegetative traits of genotypes correlate with each other, 13 tomato cultivars were subjected to long-term moderate heat (LTMH) or short-term heat shock (STHS), depending on the trait that was evaluated. LTMH caused significant decrease in performance of nearly all reproductive traits, i.e. pollen viability, pollen number, female fertility, seeded-fruit set and flower number per inflorescence, but not in inflorescence number. Considerable variation was found among cultivars, both under control and LTMH conditions. The cultivars Nagcarlang, Saladette and Malintka 101 produced a higher percentage of viable pollen under LTMH. For fruit set under LTMH condition, only cultivars that had been previously reported as being heat-tolerant produced fruits with seeds. STHS negatively affected vegetative traits concerning seedling survival and membrane stability. Correlation analysis revealed relationships between various traits within the control and heat treatments, but not between the two. Under heat stress fruit set was positively correlated with pollen viability, as well as with flower number per inflorescence. However, no significant correlations were found between vegetative and reproductive traits. Our data highlight the prominent role of pollen viability for tomato fertility under LTMH growth conditions. The observed variation in thermotolerance among different cultivars offers the possibility to decipher underlying physiological and genetic mechanisms.

We investigated the involvement of ethylene signaling in the development of the reproductive structures in tobacco ( Nicotiana tabacum L.) by studying flowers that were insensitive to ethylene. Ethylene-insensitivity was generated either by expression of the mutant etr1-1 ethylene-receptor allele from Arabidopsis thaliana or by treatment with the ethylene-perception inhibitor 1-methylcyclopropene (MCP). Development of ovaries and ovules was unaffected by ethylene-insensitivity. Anther development was also unaffected, but the final event of dehiscence was delayed and was no longer synchronous with flower opening. We showed that in these anthers degeneration of the stomium cells and dehydration were delayed. In addition, we found that MCP-treatment of detached flowers and isolated, almost mature anthers delayed dehiscence whereas ethylene-treatment accelerated dehiscence. This indicated that ethylene has a direct effect on a process that takes place in the anthers just before dehiscence. Because a similar function has been described for jasmonic acid in Arabidopsis, we suggest that ethylene acts similarly to or perhaps even in concurrence with jasmonic acid as a signaling molecule controlling the processes that lead to anther dehiscence in tobacco.