Potato Metabolic Pathways

Bringing together information on plant metabolism

Monocot Response to Drought

As an initial step towards understanding the molecular responses of monocots to drought, relevant information from refereed papers is being collated. There are clearly some similarities in the manner by which dicots and monocots respond to drought, including the involvement of carbohydrate metabolism, phosphoinositide and calcium signaling, ABA-dependent and ABA-independent pathways, and protein ubiquitination. Additionally, that some genes associated with drought response in monocots can be overexpressed in dicots (and vice versa) to increase drought resistance also seems to imply a number of signalling processes are common to both groups. However, it is important to be precise in understanding abiotic stress responses as there is potential for some processes and signaling events to be unique. Importantly, there are clear differences in the expression of many genes between different tissues (eg roots v leaves). 

Information on drought responses is being used to construct a diagram showing metabolic pathways and signalling events of drought responses in monocots. This diagram will be up-dated as new information becomes available. It is clearly not complete and only provides an indication of some of the processes involved in drought resistance. As more detailed information about drought responses is published it will be possible to provide a more complete description of events. The large number of potential signaling genes involved in the drought response pose an interesting challenge to understand how they, and their products, interact.

References on drought responses of monocots are listed at the bottom of this page.

Jiang et al. (2011) showed that over-expression of OsRIP18 (a ribosome inactivating protein gene) increased salt and drought tolerance in transgenic rice plants.

Du et al. (2011) identified an inositol 1,3,4-trisphosphate 5/6-kinase gene that is essential for drought and salt tolerance in rice.

Phung et al. (2011) showed an improved drought tolerance in rice plants expressing a protoporphyrinogen oxidase gene from Myxococcus xanthus thus demonstrating a possible role for tetrapyrroles in protecting plants from drought stress.

Campo et al. (2011) showed that a maize 14-3-3 gene ZmGF14-6 when expressed in rice conferred an increased resistance to drought though also increasing susceptibility to Fusarium verticillioides and Magnaporthe oryzae.

Saad et al. (2011) demonstrated that expressing the AlSAP gene from Aeluropus littoralis in rice increased the tolerance of rice to cold, drought and salt stresses.

Khurana et al. (2011) studied a myo-inositol-1-phosphate synthase (MIPS) gene from wheat (TaMIPS2) and showed that alternatively spliced variants from rice and Arabidopsis were involved in abiotic stresses including heat, drought, NaCl, cold, ABA, BR, SA and mannitol.

Manavalan et al. (2011) used RNAi-mediated disruption of of a rice farnesyltransferase/squalene synthase gene by maize squalene synthase to improve drought tolerance of rice.

Takeuchi et al. (2011) have shown that the root-specifc PR protein RSOsPR10 in rice is induced by drought and salt, as well as jasmonate and the ethylene precursor ACC.

Shu et al. (2011) used a protemic approach to investigate the drought response of rice. They detected 71 proteins that were affected, and using cDNA microarray and GC-MS detected 4756 differentially expressed genes and 37 differentially expressed metabolites.

Xu et al. (2011) showed increased tolerance to drought and salt stress in transgenic rice expressing ZmCBF3 with a ubiquitin promoter.

Ning et al. (2011a) demonstrated the role of the SINA E3 ubiquitin ligase OsDIS1 as a negative regulator of the drought response in rice and they suggested it may work through the posttranslational regulation of OsNEK6 (for O. sativa NIMA-related kinase 6). Ning et al. (2011b) suggested OsDIS1 interacted with OsSKIP1, a positive regulator of drought stress.

Li et al. (2011) showed that overexpression of the trehalose-6-phosphate synthase gene OsTPS1 enhanced the tolerance of rice seedlings to cold, high salinity and drought. Interestingly, the importance of trehalose-6-phosphate synthse in the response of Dicots to biotic and abiotic stress has been demonstrated by others.

Bae et al. (2011) showed that OsRDCP1, a RING domain containing protein, was induced by drought in rice. 35S:OsRDCP1 T2 transgenic rice plants showed increased tolerance to severe drought condition.

Zhang et al. (2011) showed that overexpression of a Harpin-encoding gene hrf1 in rice increased drought tolerance through ABA signaling.

Xue et al. (2011) showed that TaNAC69 is involved in adaptation of wheat to drought.

Wang et al. (2011), using microarrays, detected 5284 genes, including 251 transcription factors, that were differentially regulated in rice by drought.

Xu et al. (2011) showed that the rice gene OsMSR2 was up-regulated by cold, drought, and heat. Expression of OsMSR2 in Arabidopsis enhanced drought and salt tolerance.

Seo et al. (2011) showed that OsbHLH148 transcript levels increased with methyl jasmonate, ABA, dehydration, salt, low temperature, and wounding and that over-expression of OsbHLH148 in rice confered tolerance to drought. OsJAZ1 was possibly a transcriptional regulator of OsbHLH148.

Ouyang et al. (2011) showed that the tocopherol cyclase ortholog OsVTE1 was induced by high salt, H2O2, drought, cold, ABA and salicylic acid. Tocopherol cyclase (TC/VTE1) catalyzes the conversion of 2,3-dimethyl-5-phytyl-1,4-benzoquinone (DMPBQ) to γ-tocopherol.

Geng et al. (2011) showed that TaCPK7 from wheat responded to drought, salt, cold and H2O2.

Kumat et al. (2011) showed that OsWNK1 (a serine-threonine protein kinase 'with no lysine kinase') was up-regulated in rice in response to drought and cold

Du et al. (2010) showed that overexpression of the beta-carotene hydroxylase gene OsDSM2 increased drought tolerance in rice.

Zhou et al. (2010) described 30 microRNAs which were either up- or down-regulated by drought in rice.

Takasaki et al. (2010) showed that OsNAC5 is induced by drought, cold, high salinity, ABA and methyl jasmonate in rice and that rice plants over-expressing OsNAC5 showed an up-regulation of OsLEA3. Song et al. (2011) showed that overexpressing OsNAC5 in Arabidopsis or rice increased tolerance to drought, cold and salt. Nuruzzaman et al. (2010) have analysed the NAC transcription factor family in rice and showed that a number of them are associated with biotic or abiotic stress responses.

Hossain et al. (2010) showed that the bZIP transcription factor OsABF2 was induced by drought, salinity, cold, oxidative stress, and ABA in rice.

Zhang et al. (2010) showed that overexpression of JERF1 enhanced drought tolerance of transgenic rice by regulating the expression of OsP5CS which is involved in proline biosynthesis.

Yi et al. (2010) analysed the effectivenesss of promoters, from 6 drought inducible genes, in transgenic rice. Genes were:- Rab21, Wsi18, Lea3, Uge1, Dip1, and R1G1B.

Wang et al. (2010) investigated the regulation of auxin-related genes in Sorghum bicolor in response to abiotic stress. The genes SbIAA1, SbGH3-13, and SbLBD32 were induced by IAA, brassinosteroid, salt and drought.

Jeong et al. (2010)  showed that the root-specific expression of OsNAC10 improves drought tolerance and grain yield in rice

Cui et al. (2011) showed that over-expression  of OsDREB2A in rice increased drought tolerance.

Quan et al. (2010) showed that overexpression of tomato TSRF1 in rice enhanced drought resistance through enhanced expression of ABA putative synthesis gene SDR.

Ricachenevsky et al. (2010) showed that OsWRKY80 was up-regulated in rice by drought and was also up-regulated by Fe treatent.

Ouyang et al. (2010) showed that the expression of OsSIK1 in rice in induced by salt, drought and H2O2. Overexpression of OsSIK1 enhanced drought resistance.

Zhang et al. (2010) showed that transgenic rice expressing JERF3 were more tolerant to drought stress than non-transgenic rice. Overexpression of JERF3 in rice increased the expression of two OsP5CS genes in response to drought.

Amir Hossain et al. (2010) showed that OsABF1 was up-regulated in rice in response to drought.

Ning et al. (2010) showed that OsDSM1 (a Raf-like MAPKKK) was up-regulated in rice by drought, salt and ABA.

Feng et al. (2009) showed that the alternative oxidase AOX1a and AOX1b were up-regulated in rice seedling leaves by drought.

Kim & Kim (2009) showed that the rice transcripton factor OsAP37 was up-regulated by drought and transgenic rice using the OsCc1 promoter showed increased drought tolerance.

Zhang SW et al. (2009) described work on TLD1, a rice GH3.13 gene that encodes indole-3-acetic acid (IAA)-amido synthetase, which is suppressed in aboveground tissues under normal conditions but which is dramatically induced by drought stress.

Liu et al. (2009) showed that the rice gene OsDHODH1, encoding a putative cytosolic dihydroorotate dehydrogenase, was up-regulated by salt, drought and ABA and overexpression of OsDHODH1 in rice enhanced resistance to drought and salt.

Huang et al. (2009) showed that the zinc finger protein DST regulates drought and salt tolerance in rice  via stomatal aperture control.

Xiao et al. (2009) investigated the potential of a number of rice genes to enhance drought resistance when reintroduced into rice with various promoters. Overall, OsLOS5 and OsZAT10 showed the best results in improving drought resistance in rice when tested under field conditions.

Cheng et al. (2009) demonstrated that OsRPK1 (a leucine-rich-repeat receptor-like protein kinase) was induced by drought, cold and ABA.

Zhou et al. (2009) showed that over-expression of the wheat gene TaSTRG (salt tolerance-related gene) increased resistance of rice to drought and salt. TaSTRG is induced in wheat in response to salt, polyethylene glycol, ABA and cold.

Oh et al. (2009)  investigated 139 AP2 genes in rice and showed that overexpression of OsAP37 and OsAP59 increased tolerance to drought and salt at the vegetative stage. Importantly AP37 expressing plants showed enhanced drought tolerance in the field with increased grain yield compoared to controls.

Ding et al. (2009) showed that OsBURP05 and OsBURP16 were induced by drought, salt, cold and ABA in rice

Hou et al. (2009) showed that transgenic rice overexpressing OsSKIPa showed greater drought tolerance in the seedling and reproductive stages.

Yang et al. (2009) showed that OsMT1a (a metallothionein) was up-regulated in rice by drought and transgenic plans overrexpressing the gene showed enhanced tolerance o drought.

Kim et al. (2009) showed that rice plants produce methyl jasmonate during drought stress.

Xu et al. (2009) showed that a putative protein kinase W55a from wheat was up-regulated by drought, salt, ABA, salicylic acid, ethylene, and methyl jasmonate. Transgenic Arabidopsis overexpressing W55a exhibited a higher tolerance to drought.

Zheng et al. (2009)showed that a NAC gene, ONAC045, from rice was induced by drought, salt, low temperature, and ABA treatment in leaves and roots. Transgenic rice plants overexpressing ONAC045  showed enhanced tolerance to drought and salt.

Cai et al. (2009) showed that rice plant overexpressing the glutamine synthetase gene GS1;2 showed higher sensitivity to drought, salt and cold.

Ke et al. (2009) used proteomics to study proteins and phosphoproteins induced in drought stressed rice. They identified ten drought-responsive phosphoproteins: NAD-malate dehydrogenase, OSJNBa0084K20.14 protein, abscisic acid- and stress-inducible protein, ribosomal protein, drought-induced S-like ribonuclease, ethylene-inducible protein, guanine nucleotide-binding protein beta subunit-like protein, r40c1 protein, OSJNBb0039L24.13 protein and germin-like protein 1.

Lu et al. (2009) showed that transgenic rice overexpressing OsbZIP72 possessed enhanced drought tolerance.

Xiang et al. (2008) showed that OsbZIP23 was up-regulated by drought, salt, ABA and polyethanol glycol. Transgenic rice plants overexpressing OsbZIP23 showed improved tolerance to drought and salt.

Wu et al. (2009) showed that OsWRKY11 was induced in rice seedlings by heat shock and drought. Rice plants overexpressing OsWRKY11 showed increased tolerance to heat and drought.

Chen et al. (2008) showed that rice plants overexpressing OsDREB1G and OsDREB2B possessed improved tolerance to water stress.

Gao et al. (2008) demonstrated that rice plants overexpressing the tomato gene TERF1 showed enhanced tolerance to drought and salt.

Pasquali et al. (2008) showed that the ectopic expression of Osmyb4 in apple improved adatation to cold and drought stress.

Kumar et al. (2008) showed that MAP kinase kinase 1 from rice was regulated by drought and salt.

Huang et al. (2008) used microarray analysis to show that ZFP176 and ZFP181 (zinc finger proteins) were induced in rice by drought.

Wang et al. (2008) characterised OsDREB1F from rice and showed it was induced by salt, drought, cold and ABA. Transgenic plants expressing OsDREB1F showed enhanced tolerance to salt, drought and low temperature inboth rice and Arabidopsis.

Kobayashi et al. (2008) showed that barley HvABI5 TF was induced by drought. Expression of a wheat HvABI5 ortholog, Wabi5, was induced by low temperature, drought and ABA.

Welsch et al.(2008) showed that OsPSY3 (a phytoene synthase) was induced inrice by drought and salt, especially in roots.

Xu et al. (2008) showed that overexpresssion of OsZFP252 (a TFIIIA-type zinc finger protein) in rice enhanced tolerance to drought and salt.

Kanneganti and Gupta (2008) showed that OsiSAP8 was induced by heat, cold, salt, desiccation, submergence, wounding, heavy metals and ABA. Overexpression of the gene in both transgenic tobacco and rice conferred tolerance to salt, drought and cold stress at seed germination/seedling stage. Transgenic rice plants were tolerant to salt and drought during anthesis.

Tani et al. (2008) showed that the 12-oxophytodienoate reductase gene OsOPR7 was induced by wounding and drought.

Karaba et al. (2007) showed that expression of the Arabidopsis gene HARDY improved drought tolerance of rice.

Wu et al. (2007) described how several factors corelated with drought resistance in rice including chlorophyll content, the content of proline, the content of malondiadehyde (MDA), the activity of superoxide dismatase (SOD)and the content of peroxides (POD) and catalase (CAT).

Peres et al. (2007) showed that Orysa;EL2 mRNA levels were induced by cold, drought, and propionic acid.

Nakashima et al. (2007) showed that OsNAC6 gene expression was induced by cold, drought, salt, wounding and blast disease.

Liu et al. (2007) showed that overexpression of OsUGE-1 in Arabidopsis increased tolerance to salt, drought and freezing. Transgenic plants had a higher level of raffinose.

Liu et al. (2007) showed that OsCOIN was induced by low temperature, ABA, salt and drought. Overexpression in rice enhanced tolerance to cold, salt and droughtaccompanied by up-regulation of OsP5CS and increased proline levels.

Prashanth et al. (2008) transformed rice with a superoxide dismutase from mangrove (AmSod1) and showed that transformed plants had better tolerance to drought.

Xiang et al. (2007) surveyed CIPK genes in rice. They showed that 20 CIPK genes were differentially induced by at least one abiotic stress. Three CIPK genes (OsCIPK03, OsCIPK12, and OsCIPK15) were overexpressed in rice and the resulting transgenics showed improved tolerance to cold, drought and salt respectively.

Fu et al. (2007) showed that OsAP25  (an ERF/AP2-type TF) was induced by salt, cold, drought, ABA and ethylene.

Yang et al. (2007) studied polyamines in the drought response of rice. They showed that activities of arginine decarboxylase, S-adenosyl-L-methionine decarboxylase, and spermidine synthase in the leaves were significantly enhanced by water stress, together with an increase in putrescine, spermidine, and spermine.

Dai et al. (2007) showed that expression of OsMYB3R-2 was induced by cold, drought and salt and Arabidopsis transgenic plants overexpressing OsMYB3R-2 showed increased tolerance to cold, drought and salt.

Zhao et al. (2007) identified drought-induced microRNAs in rice including miR-169g.

Huang et al. (2007) showed that expression of OsOCPI1 (chymotrypsin inhibitor-like) was up-regulated in rice in response to drought and ABA. Overexpression of this gene in rice increased drought tolerance.

Hu et al. (2006) showed that overexpression of OsSNAC1 enhances drought resistance in transgenic rice.

Cao et al. (2006) showed that OsBIERF1, OsBIERF3 and OsBIERF4 (Oryza sativa benzothiadiazole (BTH)-induced ethylene responsive transcriptional factors) were up-regulated by salt, cold, drought and wounding.

Pramanik and Imai (2005) showed transient expression of OsTPP1 (a putative trehalose-6-phosphate phosphatase) in rice in response to cold, salt and drought.

Marè et al. (2004) showed that Hv-WRKY38 was induced by cold and drought in barley.

Yu et al. (2003) showed that OsPP2A-1 and OsPP2A-3 genes were up-regulated in rice leaves by drought.

Agrawal et al. (2002) showed that OsMSRMK2 was up-regulated by drought, wounding, protein phosphatase inhibitors, UV, heavy metals, and salt.

Kaminaka et al. (1999) showed that sodA1 Mn-SOD ) and sodCc2 (Cu/Zn-SOD ) were induced by salt and drought.

Ishitani et al. (1995) showed that mRNA levels of a betaine aldehyde dehydrogenase increase in barley in response to drought.

Qian et al. (2011) showed that a dehydrin gene Dhn6 from barley was up-regulated by drought.

Seiler et al. (2011) showed that under drought conditions barley leaves contain high concentrations of ABA and the ABA degradation products phaseic acid and diphaseic acid. They also describe regulation of genes for enzymes involved in formation and degradation of ABA glucose ester.

Pinheiro and Chaves (2011) carried out a meta analysis of drought responses and its effect on photosynthesis in Arabidopsis and baley. Amongst other comnclusions they stated that ABI1 is up-regulated by drought whilst ABI3 is usually down-regulated by drought. They also comment that ABI3 has been considered to be essential for drought recovery.

Kantar et al. (2010) identified 28 new barley miRNAs belonging to 18 distinct families. Hvu-MIR156, Hvu-MIR166, Hvu-MIR171, and Hvu-MIR408 were described as dehydration stress-responsive barley miRNAs.

Morran et al. (2011) showed that transgenic wheat and barley plants expressing TaDREB2 and TaDREB3 TFs showed greater drought tolerance relative to nontransgenic controls.

Guo et al. (2009) looked atdifferentially induced genes between drought-tolerant and drought-sensitive barley during drought stress at the reproductive stage.

Malatrasi et al. (2006) described the drought regulated gene HvBCAT-1 (a branched-chain amino acid aminotransferase).

Zhou et al. (2004) described the identification of Neophaseic acid in drought stressed barley.

Oztur et al. (2002) used microarray analysis to look at barley genes affected by drought.

Lucas et al. (2011) described TdicTMPIT1 (transmembrane protein inducible by TNF-α) which was upregulated by drought in a drought tolerant emmer wheat but not in a drought-sensitive emmer wheat.

Liu et al. (2011) described the isolation of a drought and pathogen responsive MYB TF (TaPIMP1) from wheat. Transgenic tobacco plants expressing the TaPIMP1 gene showed enhanced tolerance to drought and salt stress and also to infection by Ralstonia solanacearum.

Gao et al. (2011) showed that GmbZIP1 was induced by drought in soybean and overexpression of GmbZIP1 enhanced drought tolerance of transgenic wheat.

Li et al. (2011) showed that the expression of TaEXPB23 in wheat corresponded to water stress. Transgenic tobacco plants expressing TaEXPB23 were more drought tolerant.

Lucas et al. (2011) showed that TdicDRF1 was drought responsive in emmer wheat (Triticum turgidum ssp. dicoccoides).

Vaseva et al. (2010) observed higher ABA content, early immunodetection of dehydrins, and a significant increase of WZY2 transcript levels in drought tolerant cvs. They detected some acidic dehydrins (WCOR410b, TADHN) and showed that they were highly expressed in leaves of drought-stressed wheat. Neutral WZY2 dehydrin, TaLEA2 and TaLEA3 transcripts accumulated gradually with increasing water deficit.

Grudkowska and Zagdańska (2010) suggested that cysteine proteases are potentially involved with the drought response of wheat seedlings.

Rahaie et al. (2010) showed that TaMYBsdu1 was up-regulated by drought in wheat leaves and roots.

Li et al. (2008) described CDPK response of wheat to biotic and abiotic responses. They showed that TaCPK1, TaCPK6, TaCPK9 and TaCPK18 were regulated by drought.

Fu et al. (2009) showed that exogenous ABA and salicylic acid, salt, low temperature, dark and drought stresses can regulate the expression of TaNADP-ME1 and TaNADP-ME2 in wheat.

Gao et al. (2009) transformed wheat with a cotton DREB gene (GhDREB) and showed that the resultant transformants were more tolerant to drought, high salt, and freezing temperatures.

Wang et al. (2009) showed that when the wheat gene TaLEA3 was integrated into the grass Leymus chinensis the transgenic lines showed increased drought tolerance.

Xu et al. (2008) showed that the dehydration responsive element-binding factor TaAIDFa from wheat was responsive to drought, salinity, low temperature and ABA. Transformation of Arabidopsis with TaAIDFa  increased tolerance to drought and osmotic stress.

Jia et al. (2008) showed that the calreticulin gene TaCRT fromwheat was upregulated by drought and when transformed into tobacco (Nicotiana benthamiana) conferred increased drought tolerance.

Wang et al. (2011) showed that OsWR1 is induced by drought, ABA and salt in rice and overexpression of OsWR1 in rice increased trolerance to drought. OsWR1 is a wax synthesis regulatory gene and is a homolog of Arabidopsis WIN1/SHN1.

Kam et al. (2008) showed that expression of 37 TaZFP (zinc finger protein) genes in leaves and roots responded to drought with 74% of the drought-responsive TaZFP genes being down regulated in drought stressed roots. Sixteen of the drought-responsive TaZFP genes in leaves did not respond to ABA suggesting that some TaZFP genes are involved in ABA-independent pathways.

Kobayashi et al. (2008) showed that Wdreb2 is expressed inwheat seedlings in response to drought, cold and high salinity. They also suggested that WDREB2 regulated Wdhn13, Wrab17, Wrab18, and Wrab19 in response to stress.

Xu et al. (2007) showed that TaERF1 was upregulated in wheat in response to drought, salinity, cold, ABA, ethylene, salicylic acid and infection with Blumeria graminis f. sp. tritici.

Christov  et al. (2007) showed that DHN14 was up-regulated in wheat in response to drought and ABA.

Brini et al. (2007) showed that Arabidopsis transformed with the Triticum durum gene DHN-5 showed stronger growth under high salt concentrations or under water deprivation, and showed a faster recovery from mannitol treatment, thus contributing to improved tolerance to drought.

Hajheidarit al. (2007) used a proteomics approach to identify proteins associated with drought responses in wheat. They identified a large number of proteins which were thioredoxin targets thereby emphasising the important role of redox reactions in drought responses.

Xu et al. (2007) showed that TaPP2Ac-1 transcript levels was up-regulated in wheat (Triticum aestivum) seedlings under drought.

Rampino et al. (2006) studied the expression of dehydrins in drought stressed wheat (Triticum durum). They showed that the following genes were not expressed  in well-watered plants but were expressed in drought-stressed plants:- TdDHN9.6, TdDHN13, TdDHN15.1, TdDHN15.2 and TdDHN15.3.

Christova et al. (2006) isolated a cDNA clone for a multidomain cystatin (TaMDC1) from cold acclimated winter wheat. they showed that TaMDC1 was induced by cold, drought, salt and ABA.

Kume et al. (2005) isolated Wcbf2 (CBF/DREB1 homologs). Wcbf2 expression was induced by cold and drought but not by ABA.

Campbell et al. (2001) showed that TaHSP101B and TaHSP101C were induced in Triticum aestivum by 2hr dehydration and by ABA.

He et al. (2011) showed that TaMYB73 gene was induced by dehydration, salt and several phytohormones.

Kang et al. (2011) showed that the ribosomal gene TaL5 was upregulated in wheat by drought, salt, freezing temperatures, ABA and salicylic acid.

Ji et al. (2005) showed that the vacuolar invertase OsVIN2 was up-regulated in rice in response to drought.

Sun et al. (2011) showed that OsENAC1 was induced in rice by drought, salt, cold amnd ABA.

Zhao et al. (2011) identified 55 HD-Zip genes in the maize genome. They showed that 17 maize HD-ZIP genes were regulated by drought stress.

Jami et al. (2011) showed that rice annexin genes are regulated by various abiotic stresses including drought, salt, heat and cold. Annexins are Ca2+ dependent phospholipid-binding proteins.

Liu et al. (2012) showed that OsPFA-DSP1, a protein tyrosine phosphatase, was induced in rice in response to drought. Overexpresion of OsPFA-DSP1 in tobacco or rice increased sensitivity to drought stress suggesting it may function as a negative regulator in drought stress.

Niu et al. (2012) identified a number of stress-response WRKY genes in wheat. Transgenic Arabidopsis overexpressing TaWRKY2 showed increased tolerance to salt and drought, and overexpression of TaWRKY19 increased tolerance to salt, drought and freezing. TaWRKY2 enhanced expressions of STZ and RD29B, and TaWRKY19 activated expressions of DREB2A, RD29A, RD29B and Cor6.6.

Xia et el. (2011) showed that ZmRFP1 encodes a RING-H2 E3 ubiquitin ligase and responds to drought stress in an ABA-dependent manner in maize.

Wei et al. (2012) characterised WRKY genes in maize and showed that some of them were involved in drought responses.

Nelson et al. (2007) described a transcription factor from the nuclear factor Y family (AtNF-YB1) which was associated with drought resistance in Arabidopsis and over-expression of which enhanced drought resistance. An orthologous maize transcription factor, ZmNF-YB2, was also identified which enhanced drought resistance in maize when over-expressed.

Tang et al. (2012) showed that expression of OsbZIP46 was strongly induced in rice by drought, heat, H2O2 and ABA, but not cold or salt.

Wang et al. (2012) reported that ZmbZIP 60 from maize was upregulated by dehydration, high salinity, ABA and tunicamycin.

Zou et al. (2012) showed that transgenic rice plants overexpressing OsHsp17.0 and OsHsp23.7 had increaed tolerance to drought and salt stress compared to wild type plants.

Datta et al. (2012) examined the effect of overexpression of DREB1A from rice and Arabidopsis and DREB1B from rice and showed that rice transformed with DREB1A was more tolerant to dehydration and rice transformed with DREB1B was more salt tolerant.

Li et al. (2012) showed that HbCIPK2 expression was induced by salt, drought and ABA treatment in Hordeum brevisubulatum.

Sun et al. (2012) showed that miR156 and miR162 showed significant changes in expression levels in switchgrass (Panicum virgatum) in response to drought.

Li Y et al. (2012) investigated the rice phosphatase IBR5 and showed that drought induced WIPK activity was impaired in OsIBR5 overexpressing tobacco plants. The transgenic tobacco plants were less drought tolerant than wild type.

Nuruzzaman et al. (2012) studied expression profiles of OsNAC genes in various tissues in rice after hormone treatment and drought stress.

Mirzaei et al. (2012) studied the regulation of aquaporins, small GTPases and V-ATPases proteins in rice leaves subjected to drought stress and recovery. Most of the nine aquaporins detected were responsive to drought and nine G-proteins also increased during severe drought.

Redillas et al. (2012) showed that overexpression of OsNAC9 in rice enhanced drought resistance. Microarray experiments identified 40 up-regulated genes inroots of transgenic plants. These included 9-cis-epoxycarotenoid dioxygenase, calcium-transporting ATPase,and cinnamomyl CoA reductase. 

Bass et al. (2004) showed that the maize Rip3:2 gene (ribosome-inactivating protein ) was up-regulating by drought.

Lu et al. (2012) showed that expression of ZmSNAC1 was induced by low temperature, salinity, drought, and ABA but downregulated by salicylic acid. Overexpression of ZmSNAC1 in Arabidopsis enhanced tolerance to dehydration compared to wild-type seedlings.

Ford et al. (2011) carried out a proteomic analysis of wheat cultivars differing in drought tolerance. They noted changes consistent with an increase in oxidative stress metabolism and ROS scavenging capacity through increases in superoxide dismutases, catalases, and ROS avoidance through decreases in proteins involved in photosynthesis and the Calvin cycle.

Zhang et al. (2012) used network-based gene clustering  to study the drought respons of rice. They identified 2,607 rice genes that showed significant changes in gene expression under drought stress, and they identified 15 gene nodules.

Ali and Komatsu (2006) used a proteomic approach to investigate drought induced changes in rice leaf sheath.

Sato and Yokoya (2008) showed that overexpression of a small heat shock protein sHSP17.7 in rice increased drought tolerance.

Yeap et al. (2012) showed that EgRBP42, encoding a member of the plant heterogeneous nuclear ribonucleoprotein (hnRNP)-like RBP family from oil palm (Elaeis guineensis Jacq.) was up-regulated by salinity, drought, submergence, cold and heat stresses in leaf discs.

Omidvar et al. (2012) showed that in oil palm the genes EABF and EABF1 were upregulated in response to ABA, ethylene, methyl jasmonate, drought, cold and high-salinity treatments.

Singh et al. (2012) provided detailed microarray based expression analysis and expression profiles of PLD genes in rice, under three abiotic stresses (salt, cold and drought) and different developmental stages (3-vegetative stages and 11-reproductive stages).

Peng et al. (2012) studied the CCCH-type zinc finger family in maize. Seven of these genes showed differential expression patterns among five representative maize tissues and over time in response to abscisic acid and drought treatments.

Chen et al. (2012) showed that OsbZIP16 was induced in rice in response to drought.  Transgenic rice plants overexpressing OsbZIP16 exhibited significantly improved drought resistance.

Peng et al. (2010) showed that PLDalpha1 from foxtail millet (Setaria italica) is up-regulated by dehydration, ABA and NaCl . Overexpression of SiPLDalpha1 in Arabidopsis enhanced tolerance to drought stress.

Ramegowda et al. (2012) showed that EcNAC1 was upregulated in finger millet by water deficit and salt.

Barrera-Figueroa et al. (2012) identified 18, 15, and 13 miRNAs that were regulated by drought, cold and salt stress conditions, respectively, in rice inflorescences.

Zhang et al. (2012) showed that overexpression of OsPIN3t in rice increased drought tolerance.

Sreedharan et al. (2012) showed that MusaSAP1 was up-regulated by drought, salt, cold, heat, oxidative stress and ABA.

Begcy et al. (2012) showed that sugarcane drought responsive 1 (Scdr1) was upregulated by drought and overexpression of Scdr1 in transgenic tobacco plants increased their tolerance to drought, salinity and oxidative stress.

Wendelboe-Nelson and Morris (2012) carried out a protemic comparison of barley cv Golden Promise (drought susceptible) and cv Basrah (greater drought tolerance). The variety Basrah was characterised by constitutive expression or higher drought-induced expression levels of proteins regulating ROS production and protein folding.

Duan et al. (2012) showed that OsMIOX from upland rice (Oryza sativa L. cv. IRAT109) was expressed predominantly in the roots and induced by drought, H2O2, salt, cold and abscisic acid. The survival rate of leaves from the transgenic rice lines was higher than that of the wild type plants under polyethylene glycol treatment.

Xiao et al. (2012) showed that expression of the TaHPS gene in Triticum aestivum  was induced by abscisic acid, salt and drought. Overexpression of TaHPS in Arabidopsis increased tolerance to salt and drought.

Kim et al. (2003) showed that OsEDR1 (a MAPKKK) has a constitutive expression in rice seedling leaves and is further up-regulated by wounding, jasmonic acid, salicylic acid, ethylene (ethephon), ABA, hydrogen peroxide, protein phosphatase inhibitors, chitosan, drought, high salt and sugar, and heavy metals.

Singh and Ghosh (2012) showed that OsGS2 and OsGS1;1 expression may contribute to drought tolerance in a drought resistant cultivar.

Jeong et al. (2012) showed that overexpression of OsNAC5 in rice roots enlarges roots and enhances drought tolerance and grain yield under field conditions.

Zhang et al. (2012) showed that expression of Oshox22 (an HD-Zip gene) is strongly induced by salt, ABA, and PEG, and weakly by cold stress. Transgenic rice over-expressing Oshox22 showed increased sensitivity to ABA, increased ABA content, and decreased drought and salt tolerances.

Du et al. (2012) showed that expression of OsGH3-2 in rice was induced by drought but was suppressed by cold.

You et al. (2012) showed OsOAT is a direct target of SNAC2 and overexpression of OsOAT enhanced drought and osmotic stress tolerance. They claimed that both ABA-dependent and ABA-independent pathways contributed to the drought-induced expression of OsOAT.

Gentile et al. (2012) identified 13 mature miRNAs that were differentially expressed in drought-stressed field-grown sugarcane plants

Ashoub et al. (2012)  identified differentially expressed proteins in barley land races (differing in drought response) in response to drought.

Liu et al. (2012) showed that drought stress tolerance of ZmPIS (phosphatidylinositol synthase) sense transgenic plants were enhanced at the pre-flowering stages compared to WT maize plants.

Tang et al. (2012) showed that expression of the AtbZIP60 gene enhanced salt, drought, and cold tolerance in rice and white pine transgenic cell lines.

Zhou et al. (2012) showed that expression of  the aquaporin gene TaAQP7 in wheat was induced by dehydration, PEG, ABA, and H2O2. Overexpression in tobacco enhanced drought tolerance. 

Zhou et al. (2013) showed that transgenic creeping bentgrass (Agrostis stolonifera) overexpressing a rice miR319 gene showed enhanced drought and salt tolerance.

Asad et al. (2013) showed that expression of OsTZF1 was induced in rice by drought, salt, H2O2, ABA, methyl jasmonate, and salicylic acid.

Yadav et al. (2012) showed that RGA1(I) (a Gα subunit) was upregulated by NaCl, cold and drought stress and down regulated by elevated temperature.

Lu et al. (2013) showed that overexpression of Arabidopsis molybdenum cofactor sulfurase gene (LOS5) gene in maize enhanced the expression of ZmAO and aldehyde oxidase activity, leading to increased drought tolerance.

Liu et al. (2013) showed that transgenic rice seedlings over-expressing OsHsfA7 were more tolerant to drought and salt.

Manmathan et al. (2013) showed that Era1 (enhanced response to abscisic acid) and Sal1 (inositol polyphosphate 1-phosphatase) play important roles in conferring drought tolerance in wheat.

de Abreu-Neto et al. (2013) showed that the rice gene OsHIPP41 is highly expressed in response to cold and drought stresses.

Cal et al. (2013) used transcriptome profiling of the leaf elongation zone of contrasting rice cultivars under drought conditions.

Smita et al. (2013) identified a number of drought responsive gene in rice and suggested that drought stress mediated upregulated gene expression which was coordinated through an ABA-dependent signaling pathway across tissues.

Saad et al. (2013) showed that Chinese wheat variety Yangmai12 expressing the rice NAC1 gene (SNAC1) showed enhanced tolerance to drought and salinity.

Huang et al. (2013) showed that expression of OsERF3 was induced by drought, salt, ACC and ABA and that the EAR motif is required for OsERF3 to transcriptionally regulate the ethylene synthesis and drought tolerance in rice.

Luang et al (2013) showed that Os9BGlu31 (a glycoside hydrolase family GH1 transglycosidase) is upregulated in rice by drought, ABA, ethephon, methyl jasmonate, 2,4-D and kinetin.

Verlotta et al. (2013) showed that there are inducible secretory phospholipases (sPLA isoforms) in durum wheat that are orchestrating the response to drought.

Zhang et al. (2013) showed that the cytosolic ascorbate peroxidase OsAPX2 plays an important part in protecting rice from drought, salt and cold stresses.

Vilela et al. (2013) showed that ZmSNAC1 is a substrate of ZmOST1. The kinase OPEN STOMATA 1 (OST1) plays an important role in regulating drought stress signaling, particularly stomatal closure.

Park et al. (2013) showed that a rice immunophilin gene OsFKBP16-3 was induced by salt, drought, high light, hydrogene peroxide, heat and methyl viologen, and transgenic Arabidopsis and rice constitutively expressing OsFKBP16-3 showed increase tolerance to salinity, drought, and oxidative stress.

Tian et al. (2013) showed that overexpression of TaSnRK2.3 in Arabidopsis resulted in an improved root system and significantly enhanced tolerance to drought, salt, and freezing stresses.

Joo et al. (2013) showed that the expression of both OsASR1 and OsASR3 was induced by drought and that overexpression of either OsASR1 or OsASR3 in transgenic rice plants increased their tolerance to drought and cold stress. 

Wang et al. (2013) showed that drought stress substantially upregulated the expression of OsNox1-3, OsNox5, OsNox9, and OsFRO1, but downregulated OsNox6 in rice.

Guo et al. (2013) showed that OsDIL was primarily expressed in the anther and mainly responsive to abiotic stresses, including drought, cold, NaCl, and ABA. Compared with wild type, the OsDIL-overexpressing transgenic rice plants were more tolerant to drought stress during vegetative development and showed less severe tapetal defects and fewer defective anther sacs when treated with drought at the reproductive stage.

Hadiarto and Tran (2011) provide an excellent review of drought responsive genes in rice.

Xu et al. (203) showed that expression of OsSDS1 was upregulated by cold, drought and heat. Expression of OsSDS1 conferred decreased tolerance to salt and drought in Arabidopsis suggesting that OsSDS1 may act as a negative regulator of salt and drought tolerance in plants.

Yu et al. (2013) showed that transgenic rice plants expressing the Arabidopsis HD-ZIP transcription factor EDT1/HDG11 showed increased drought tolerance.

Xia et al. (2013) showed that transcript levels of ZmSKD1 were upregulated by salt or drought stress and overexpression of ZmSKD1 in tobacco increased tolerance tosalt and drought.

Liu et al. (2013) showed that OsbZIP71 was expressed in rice by drought, PEG, and ABA. Overexpression of bZIP71 increased tolerance to drought, salt and PEG. OsVHA-B, OsNHX1, COR413-TM1, and OsMyb4, were up-regulated in overexpressing lines.

Xiang et al. (2013) showed that OsHsfB2b negatively regulates salt and drought tolerance in rice.

Anna et al. (2013) suggested that progesterone is a part of the wheat response to stress factors (drought).

Nakashima et al. (2013) compared drought-responsive promoters in transgenic rice. They showed that transgenic rice plants overexpressing OsNAC6 under the control of the Oshox24 promoter showed increased tolerance to drought and high salinity.

Ham et al. (2013) showed that OsSHSP1 (Os03g16030) and OsSHSP2 (Os01g04380) were induced by salt, drought, ABA and slicylic acid but not cold, jasmonic acid, or ethylene.

Zhu and Xiong (2013) showed that DWA1 was stronly induced in rice by drought. DWA1 may regulate drought-induced cuticular wax deposition.

Huda et al. (2013) show that OsACA6 (a Ca2+ ATPase ) is upregulated by drought, salt, ABA and heat.

Xiao et al. (2013) suggested that OsWRKY13 regulates the antagonistic crosstalk between drought and disease resistance pathways by directly suppressing OsSNAC1 and OsWRKY45-1 in rice.

Joo et al. (2013) showed that over-expression of BvMTSH in rice enhances drought-stress tolerance.

Lim et al. (2013) showed that plants overexpressing the RING E3 ligase OSCTR1 showed improvement in their tolerance against severe water deficits.

Yang et al. (2014) showed that AtGRP2- or AtGRP7-expressing transgenic rice plants showed much higher recovery rates and grain yields compared with the wild-type plants under drought stress conditions.

Fang et al. (2013) showed that drought stress caused a significant increase in the expression of four histone acetyltransferases (HATs )(OsHAC703, OsHAG703, OsHAF701 and OsHAM701) in rice plants.

Degenkolbe et al. (2013) published a very nice paper on drought tolerance markers in rice by expression and metabolite profiling.

Feng et al. (2014) suggest that the conserved miR164-targeted NAC genes may be negative regulators of drought tolerance in rice.

Xu et al. (2014) showed that heterologous expression of banana MaPIP1;1 in Arabidopsis confers salt and drought stress tolerance.

Reis et al. (2014) showed that induced expression of AtDREB2A CA in sugarcane enhanced its drought tolerance.

Xiong et al. (2014) showed that overexpression of OsMYB48-1 in rice improved tolerance to simulated drought and salinity stress caused by mannitol, PEG and NaCl, and drought stress in soil. 

Okay et al. (2014) looked at a number of wheat WRKY transcription factors in response to drought and showed differential expression of TaWRKY16, TaWRKY24, TaWRKY59, TaWRKY61 and TaWRKY82 genes.

Wei et al. (2014) showed that OsCPK9 positively regulates drought stress tolerance and spikelet fertility. Overexpression of OsCPK9 improved drought tolerance.

Nguyen  et al. (2014) suggested a gene network involving 9 rice ABC transporters are differentially regulated by drought stress in roots. 

Wang et al. (2014) identified 303 maize microRNAs expressed under drought stress and showed that some played critical roles in drought responsive signaling pathways. 

Akdogan et al. (2015) investigated drought-responsive miRNAs in the root and leaf of bread wheat. 

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