This report provides direct evidence that strigolactone (SL) positively regulates drought and high salinity responses in Arabidopsis. Both SL-deficient and SL-response [more axillary growth (max)] mutants exhibited hypersensitivity to drought and salt stress, which was associated with shoot- rather than root-related traits. Exogenous SL treatment rescued the drought-sensitive phenotype of the SL-deficient mutants but not of the SL-response mutant, and enhanced drought tolerance of WT plants, confirming the role of SL as a positive regulator in stress response. In agreement with the drought-sensitive phenotype, max mutants exhibited increased leaf stomatal density relative to WT and slower abscisic acid (ABA)-induced stomatal closure. Compared with WT, the max mutants exhibited increased leaf water loss rate during dehydration and decreased ABA responsiveness during germination and postgermination. Collectively, these results indicate that cross-talk between SL and ABA plays an important role in integrating stress signals to regulate stomatal development and function. Additionally, a comparative microarray analysis of the leaves of the SL-response max2 mutant and WT plants under normal and dehydrative conditions revealed an SL-mediated network controlling plant responses to stress via many stress- and/or ABA-responsive and cytokinin metabolism-related genes. Our results demonstrate that plants integrate multiple hormone-response pathways for adaptation to environmental stress. Based on our results, genetic modulation of SL content/response could be applied as a potential approach to reduce the negative impact of abiotic stress on crop productivity.

https://www.pnas.org/content/pnas/111/2/851.full.pdf

Positive regulatory role of strigolactone in plant responses to drought and salt stress

Drought stress is a severe environmental constraint among the emerging problems. Plants are highly vulnerable to drought stress and a severe decrease in yield was recorded in the last few decades. So, it is highly desirable to understand the mechanism of drought tolerance in plants and consequently enhance the tolerance against drought stress. Phytohormones are known to play vital roles in regulating various phenomenons in plants to acclimatize to varying drought environment. Abscisic acid (ABA) is considered the main hormone which intensifies drought tolerance in plants through various morpho-physiological and molecular processes including stomata regulation, root development, and initiation of ABA-dependent pathway. In addition, jasmonic acid (JA), salicylic acid (SA) ethylene (ET), auxins (IAA), gibberellins (GAs), cytokinins (CKs), and brassinosteroids (BRs) are also very important phytohormones to congregate the challenges of drought stress. However, these hormones are usually cross talk with each other to increase the survival of plants in drought conditions. On the other hand, the transgenic approach is currently the most accepted technique to engineer the genes responsible for the synthesis of phytohormones in drought stress response. Our present review highlights the regulatory circuits of phytohormones in drought tolerance mechanism.

Phytohormones enhanced drought tolerance in plants: a coping strategy.

The phytohormone cytokinin was originally discovered as a regulator of cell division. Later, it was described to be involved in regulating numerous processes in plant growth and development including meristem activity, tissue patterning, and organ size. More recently, diverse functions for cytokinin in the response to abiotic and biotic stresses have been reported. Cytokinin is required for the defence against high light stress and to protect plants from a novel type of abiotic stress caused by an altered photoperiod. Additionally, cytokinin has a role in the response to temperature, drought, osmotic, salt, and nutrient stress. Similarly, the full response to certain plant pathogens and herbivores requires a functional cytokinin signalling pathway. Conversely, different types of stress impact cytokinin homeostasis. The diverse functions of cytokinin in responses to stress and crosstalk with other hormones are described. Its emerging roles as a priming agent and as a regulator of growth-defence trade-offs are discussed.

Cytokinin action in response to abiotic and biotic stresses in plants.

Abiotic stresses due to environmental factors could adversely affect the growth and development of crops. Among the abiotic stresses, drought and heat stress are two critical threats to crop growth and sustainable agriculture worldwide. Considering global climate change, incidence of combined drought and heat stress is likely to increase. The aim of this study was to shed light on plant growth performance and leaf physiology of three tomatoes cultivars (‘Arvento’, ‘LA1994’ and ‘LA2093’) under control, drought, heat and combined stress. Shoot fresh and dry weight, leaf area and relative water content of all cultivars significantly decreased under drought and combined stress as compared to control. The net photosynthesis and starch content were significantly lower under drought and combined stress than control in the three cultivars. Stomata and pore length of the three cultivars significantly decreased under drought and combined stress as compared to control. The tomato ‘Arvento’ was more affected by heat stress than ‘LA1994’ and ‘LA2093’ due to significant decreases in shoot dry weight, chlorophyll a and carotenoid content, starch content and NPQ (non-photochemical quenching) only in ‘Arvento’ under heat treatment. By comparison, the two heat-tolerant tomatoes were more affected by drought stress compared to ‘Arvento’ as shown by small stomatal and pore area, decreased sucrose content, ΦPSII (quantum yield of photosystem II), ETR (electron transport rate) and qL (fraction of open PSII centers) in ‘LA1994’ and ‘LA2093’. The three cultivars showed similar response when subjected to the combination of drought and heat stress as shown by most physiological parameters, even though only ‘LA1994’ and ‘LA2093’ showed decreased Fv/Fm (maximum potential quantum efficiency of photosystem II), ΦPSII, ETR and qL under combined stress. The cultivars differing in heat sensitivity did not show difference in the combined stress sensitivity, indicating that selection for tomatoes with combined stress tolerance might not be correlated with the single stress tolerance. In this study, drought stress had a predominant effect on tomato over heat stress, which explained why simultaneous application of heat and drought revealed similar physiological responses to the drought stress. These results will uncover the difference and linkage between the physiological response of tomatoes to drought, heat and combined stress and be important for the selection and breeding of tolerant tomato cultivars under single and combine stress.

/pdf/drought-stress-had-a-predominant-effect-over-heat-stress-on-3l0ju7wqot.pdf

Drought stress had a predominant effect over heat stress on three tomato cultivars subjected to combined stress.

Plants have evolved elaborate mechanisms for sensing and responding to sub-optimal environmental conditions. Abiotic stresses caused by these conditions trigger a wide range of local and long-distance signals which must be co-ordinated and integrated into whole-plant processes, such as development, in order for the plant to respond properly and survive. Several hormones function as key regulators of stress tolerance, connecting local stimuli to systemic responses. Cytokinin is a hormone well known for its role in numerous aspects of growth and development, although abundant evidence also indicates that cytokinin functions in stress responses as well. At present, a full understanding of the effects of cytokinin on plant resistance to stress is lacking, possibly as a result of the complex interactions between cytokinin and stress signalling. Current knowledge of the physiological relationship between cytokinin and abiotic stress, based on measurements of cytokinin levels under stress conditions and the effects of cytokinin treatment on stress tolerance, has been examined here. A pattern of transcriptional regulation of stress-related genes by cytokinin in different plant species has also been identified. In addition, research regarding the role of specific cytokinin signalling components in a variety of stress responses is presented. We discuss what this body of research collectively implies with regard to cross-talk between cytokinin and abiotic stress tolerance.

/pdf/interactions-between-cytokinin-signalling-and-abiotic-stress-oq5vkl7ksa.pdf

Interactions between cytokinin signalling and abiotic stress responses

Responses to drought, heat, and combined stress were compared in tobacco (Nicotiana tabacum L.) plants ectopically expressing the cytokinin oxidase/dehydrogenase CKX1 gene of Arabidopsis thaliana L. under the control of either the predominantly root-expressed WRKY6 promoter or the constitutive 35S promoter, and in the wild type. WRKY6:CKX1 plants exhibited high CKX activity in the roots under control conditions. Under stress, the activity of the WRKY6 promoter was down-regulated and the concomitantly reduced cytokinin degradation coincided with raised bioactive cytokinin levels during the early phase of the stress response, which might contribute to enhanced stress tolerance of this genotype. Constitutive expression of CKX1 resulted in an enlarged root system, a stunted, dwarf shoot phenotype, and a low basal level of expression of the dehydration marker gene ERD10B. The high drought tolerance of this genotype was associated with a relatively moderate drop in leaf water potential and a significant decrease in leaf osmotic potential. Basal expression of the proline biosynthetic gene P5CSA was raised. Both wild-type and WRKY6:CKX1 plants responded to heat stress by transient elevation of stomatal conductance, which correlated with an enhanced abscisic acid catabolism. 35S:CKX1 transgenic plants exhibited a small and delayed stomatal response. Nevertheless, they maintained a lower leaf temperature than the other genotypes. Heat shock applied to drought-stressed plants exaggerated the negative stress effects, probably due to the additional water loss caused by a transient stimulation of transpiration. The results indicate that modulation of cytokinin levels may positively affect plant responses to abiotic stress through a variety of physiological mechanisms.

/pdf/enhanced-drought-and-heat-stress-tolerance-of-tobacco-plants-3razf5d8qt.pdf

Enhanced drought and heat stress tolerance of tobacco plants with ectopically enhanced cytokinin oxidase/dehydrogenase gene expression

Cytokinin oxidase/dehydrogenase overexpression modifies antioxidant defense against heat, drought and their combination in Nicotiana tabacum plants

The responses of antioxidant enzymes (AOE) ascorbate peroxidase (APX), glutathione reductase (GR), superoxide dismutase (SOD), and catalase (CAT) in soluble protein extracts from leaves and roots of tobacco (Nicotiana tabacum L. cv. Samsun NN) plants to the drought stress, salinity and enhanced zinc concentration were investigated. The studied tobacco included wild-type (WT) and transgenic plants (AtCKX2) harbouring the cytokinin oxidase/dehydrogenase gene under control of 35S promoter from Arabidopsis thaliana (AtCKX2). The transgenic plants exhibited highly enhanced CKX activity and decreased contents of cytokinins and abscisic acid in both leaves and roots, altered phenotype, retarded growth, and postponed senescence onset. Under control conditions, the AtCKX2 plants exhibited noticeably higher activity of GR in leaves and APX and SOD in roots. CAT activity in leaves always decreased upon stresses in WT while increased in AtCKX2 plants. On the contrary, the SOD activity was enhanced in WT but declined in AtCKX2 leaves. In roots, the APX activity prevailingly increased in WT while mainly decreased in AtCKX2 in response to the stresses. Both WT and AtCKX2 leaves as well as roots exhibited elevated abscisic acid content and increased CKX activity under all stresses while endogenous CKs and IAA contents were not much affected by stress treatments in either WT or transgenic plants.

https://link.springer.com/content/pdf/10.1007%2Fs10535-010-0082-3.pdf

Effect of abiotic stresses on the activity of antioxidative enzymes and contents of phytohormones in wild type and AtCKX2 transgenic tobacco plants.

Plant ageing and senescence are associated with increased levels of reactive oxygen species. Level of cytokinins, the apparent inhibitors of plant senescence, is controlled by their irreversible degradation catalysed by cytokinin oxidase/dehydrogenase (CKX). We investigated the CKX activity, cytokinin concentration, and activities of antioxidative enzymes in tobacco (Nicotiana tabacum L. cv. Samsun NN) overexpressing the Arabidopsis gene for AtCKX2, targeted for extracellular secretion pathway. The control and AtCKX2 plants differed substantially in their phenotypes. When the lowest leaves in controls became yellow all leaves in AtCKX2 tobacco still remained green. Activities of antioxidant enzymes decreased with leaf age in both tobacco plants except for ascorbate peroxidase (APX) in the old leaves and glutathione reductase (GR) in young leaves. Enhancement of GR activity at all leaf stages, an increase of superoxide dismutase and a decline of catalase in young leaves, as well as an increase of APX in the oldest leaves were observed in AtCKX2 plant compared to control. Similar changes were detected after determination of isoenzymes on zymograms. It is evident that AtCKX2 plants had postponed onset of senescence despite the significantly lowered level of cytokinins. Enhanced antioxidant protection, especially in the oldest leaves, could subsidise this phenomenon.

Zuzana Lubovská

Papers

Enhanced drought and heat stress tolerance of tobacco plants with ectopically enhanced cytokinin oxidase/dehydrogenase gene expression

Cytokinin oxidase/dehydrogenase overexpression modifies antioxidant defense against heat, drought and their combination in Nicotiana tabacum plants

Effect of abiotic stresses on the activity of antioxidative enzymes and contents of phytohormones in wild type and AtCKX2 transgenic tobacco plants.

Antioxidant enzymatic protection during tobacco leaf ageing is affected by cytokinin depletion