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...l., 2010). For instance, constitutive overexpression of Arabidopsis thaliana DREB1A or rice DREB1A confers freezing and dehydration tolerance to Arabidopsis, but results in severe growth retardation (=-=Kasuga et al., 1999-=-; Liu et al., 1998; Dubouzet et al., 2003). The constitutive expression of a pigeonpea proline-rich protein in Arabidopsis confers tolerance to osmotic, salt, and heat stress, but produces plants that...

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...ieterse et al., 2009). It acts as a key repressor of the SA pathway (Laurie-Berry et al., 2006). MYC2 has also been found to be activated by ABA, and myc2 mutants lack ABA-responsive gene expression (=-=Abe et al., 2003-=-). Therefore MYC2 may act as a centralregulator by which ABA controls biotic stress signalling pathways (Anderson et al., 2004; Asselbergh et al., 2008b; Pieterse et al., 2009). This partial synergy b...

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...Zhang et al., 2006). During biotic stress, plant transmembrane receptors such as FLS2 detect PAMPs and trigger MAPK cascades in order to establish pathogen response signalling (Nakagami et al., 2004; =-=Chinchilla et al., 2007-=-). During abiotic stress, MAPK cascades such as the MEKK1/ MKK2/MPK4/MPK6 pathway are induced, for instance following activation of the transmembrane osmoreceptor Table 1. Continued Gene Type Species ...

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..., and abiotic and biotic stress. It induces genes such as ascorbate peroxidase (APX1), and when overexpressed conferred tolerance to oxidative stress, freezing, and high light (Rizhsky et al., 2004a; =-=Davletova et al., 2005-=-; Vogel et al., 2005; Fujita et al., 2006). Similarly, ZAT7 is activated in response to oxidative stress, and confers tolerance to salt, cold, and oxidative stress when expressed ectopically (Rizhsky ...

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...-transcriptionally or through phosphorylation (Herrgard et al., 2003). In addition, correlation between proteomic data and transcript abundance measured by microarrays is surprisingly low (R 0.24) (=-=Fu et al., 2009-=-). In order to obtain a more accurate description of components in stress signalling pathways, metabolic and proteomic analyses are becoming increasingly utilized (Koussevitzky et al., 2008; Shulaev e...

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...rulent P. syringae strain (Mohr and Cahill, 2003), in tomato to B. cinerea and Erwinia chrysanthemi (Audenaert et al., 2002; Asselbergh et al., 2008a), in rice to the blast fungus Magnaportha grisea (=-=Koga et al., 2004-=-), and in potato to the pathogens Phytophthora infestans and Cladosporium cucumerinum (Henfling et al., 1980). In accordance with this, a lack of ABA can induce a higher level of pathogen Biotic and a...

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...ngly low (R 0.24) (Fu et al., 2009). In order to obtain a more accurate description of components in stress signalling pathways, metabolic and proteomic analyses are becoming increasingly utilized (=-=Koussevitzky et al., 2008-=-; Shulaev et al., 2008; Urano et al., 2009; Hofmann et al., 2010). The integration of these data with transcriptome results will provide the most powerful tool for characterizing such complex plant pr...

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... 2005; Fujita et al., 2006). Similarly, ZAT7 is activated in response to oxidative stress, and confers tolerance to salt, cold, and oxidative stress when expressed ectopically (Rizhsky et al., 2004a; =-=Ciftci-Yilmaz et al., 2007-=-). ZAT7 also induces the expression of various defence transcripts, includingWRKY70, a gene that modulates cross-talk between the SA and JA pathogen response pathways (Li et al., 2006; Ciftci-Yilmaz e...

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... Khan and Khan, 1996; EnglishLoeb et al., 1997; Smit and Vamerali, 1998; Audebert et al., 2000), whereas long-term abiotic stress can weaken plant defences and cause enhanced pathogen susceptibility (=-=Amtmann et al., 2008-=-; Goel et al., 2008; Mittler and Blumwald, 2010). Despite this, few reports in the literature have focused on understanding the interaction between biotic and abiotic stresses. A range of case studies...

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... biotic and abiotic stresses has been achieved. In maize, breeding programmes have led to plants which are tolerant to drought and have additional resistance to the parasitic weed Striga hermonthica (=-=Banziger et al., 2006-=-; Badu-Apraku and Yallou, 2009). However, research programmes aimed at developing tolerance to a particular stress do not necessarily test susceptibility to other biotic or abiotic stresses. This can ...

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...o multiple abiotic or multiple biotic stresses and provided targets for the improvement of stress tolerance (Voelckel and Baldwin, 2004; Hewezi et al., 2008; Priyanka et al., 2010; Wang et al., 2010; =-=Grigorova et al., 2011-=-). A combination of high light and high temperature activated a specific subset of differentially expressed genes in sunflower (Hewezi et al., 2008). In wheat, simultaneous drought and heat stress act...

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...can cause detrimental effects to plant pathogen resistance. In both sorghum and the common bean, drought-treated plants had a higher susceptibility to the charcoal rot fungus Macrophomina phaseolina (=-=Diourte et al., 1995-=-; Mayek-Perez et al., 2002). Similarly, drought stress increased the spread of fungal and bacterial leaf scorch symptoms on date palms and the Parthenocissus quinquefolia vine (McElrone et al., 2001; ...

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...2006; Zhang et al., 2006; AbuQamar et al., 2009; Dubos et al., 2010), and that secondary metabolites such as flavonoids accumulate in response to and provide protection from a wide range of stresses (=-=EnglishLoeb et al., 1997-=-; Treutter, 2006; Cle et al., 2008; Atkinson et al., 2011). The occurrence of simultaneous biotic and abiotic stresses presents an added degree of complexity, as the responses to these are largely con...

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...on between growth and defence against stress, as responding to stress can be costly and reduce fitness in terms of growth and yield (Herms and Mattson, 1992; Tian et al., 2003; Smith and Stitt, 2007; =-=Bechtold et al., 2010-=-). This cost is highlighted by examples where the experimental transgenic expression of a stressresponsive gene confers stress tolerance but to the detriment of growth and yield (Bechtold et al., 2010...

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...cago truncatula the drought-induced production of MtDCL1 and MtAGO1, enzymes that regulate the production and processing of miRNAs, provides a mechanism for the regulation of these small RNA species (=-=Capitao et al., 2011-=-). Improvement of stress tolerance by manipulation of miRNA regulation has also been demonstrated. Transgenic Arabidopsis plants expressing a form of the chloroplastic superoxide dismutase CSD2 that i...

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...the whitefly pest Bemisia tabaci (Otim et al., 2006); new drought-tolerant aerobic rice varieties that were particularly susceptible to nematode infection, perhaps due to their increased root length (=-=Kreye et al., 2009-=-); and transgenic cotton plants expressing the Bacillus thuringiensis (Bt) insecticidal protein Cry that showed a reduction in the level of the protein during periods of high temperature, elevated CO2...

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...enaert et al., 2002; Asselbergh et al., 2008a), in rice to the blast fungus Magnaportha grisea (Koga et al., 2004), and in potato to the pathogens Phytophthora infestans and Cladosporium cucumerinum (=-=Henfling et al., 1980-=-). In accordance with this, a lack of ABA can induce a higher level of pathogen Biotic and abiotic stress interaction | 5 of 21sat Pennsylvania State U niversity on M arch 6, 2016 http://jxb.oxfordjou...

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..., stabilizing soil structure, increasing osmolyte production, and improving abscisic acid (ABA) regulation, amongst others (Alami et al., 2000; Saravanakumar and Samiyappan, 2007; Aroca et al., 2008; =-=Kohler et al., 2008-=-; Grover et al., 2011). The employment of such microbes may provide an alternative strategy to genetic engineering and plant breeding for delivering swift but effective increases in crop stress tolera...