Regulon: Uma visão geral do sistema regulador da transcrição do estresse abiótico em plantas e papel nas plantas transgênica

Warsi, M. K.; Howladar, S. M.; Alsharif, M. A.

M. K. Warsi ^**e-mail: mwarsi@uj.edu.sa

Department of Biochemistry, College of Science, University of Jeddah, Jeddah, Saudi Arabia

http://orcid.org/0000-0002-4860-8360

S. M. Howladar

Department of Biology, College of Science, University of Jeddah, Jeddah, Saudi Arabia

M. A. Alsharif

Architecture Department, Faculty of Engineering. Albaha University, Albaha, Saudi Arabia

*e-mail: mwarsi@uj.edu.sa

Figuras (5)
Tabelas (2)

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Figure 5
DREB gene family express under drought and sat stress condition in different plants. GmDREBa, GmDREBb, GmDREBc (Li et al., 2005LI, X.P., TIAN, A.G., LUO, G.Z., GONG, Z.Z., ZHANG, J.S. and CHEN, S.Y., 2005. Soybean DRE-binding transcription factors that are responsive to abiotic stresses. Theoretical and Applied Genetics, vol. 110, no. 8, pp. 1355-1362. http://dx.doi.org/10.1007/s00122-004-1867-6. PMid:15841365.
http://dx.doi.org/10.1007/s00122-004-186... ), GmDREBa (Chen et al., 2007CHEN, M., WANG, Q.Y., CHENG, X.G., XU, Z.S., LI, L.C., YE, X.G., XIA, L.Q. and MA, Y.Z., 2007. GmDREB2, a soybean DRE-binding transcription factor, conferred drought and high-salt tolerance in transgenic plants. Biochemical and Biophysical Research Communications, vol. 353, no. 2, pp. 299-305. http://dx.doi.org/10.1016/j.bbrc.2006.12.027. PMid:17178106.
http://dx.doi.org/10.1016/j.bbrc.2006.12... ), CAP2 (Shukla et al., 2006SHUKLA, R.K., RAHA, S., TRIPATHI, V. and CHATTOPADHYAY, D., 2006. Expression of CAP2, an APETALA2-family transcription factor from chickpea, enhances growth and tolerance to dehydration and salt stress in transgenic tobacco. Plant Physiology, vol. 142, no. 1, pp. 113-123. http://dx.doi.org/10.1104/pp.106.081752. PMid:16844836.
http://dx.doi.org/10.1104/pp.106.081752... ), DREB2A (Liu et al., 1998LIU, Q., KASUGA, M., SAKUMA, Y., ABE, H., MIURA, S., YAMAGUCHI-SHINOZAKI, K. and SHINOZAKI, K., 1998. Two transcription factors, DREB1 and DREB2, with an EREBP/AP2 DNA binding domain separate two cellular signal transduction pathways in drought-and low-temperature-responsive gene expression, respectively, in Arabidopsis. The Plant Cell, vol. 10, no. 8, pp. 1391-1406. http://dx.doi.org/10.1105/tpc.10.8.1391. PMid:9707537.
http://dx.doi.org/10.1105/tpc.10.8.1391... ), OsDREB1C & OsDREB2A (Dubouzet et al., 2003DUBOUZET, J.G., SAKUMA, Y., ITO, Y., KASUGA, M., DUBOUZET, E.G., MIURA, S., SEKI, M., SHINOZAKI, K. and YAMAGUCHI-SHINOZAKI, K., 2003. OsDREB genes in rice, Oryza sativa L., encode transcription activators that function in drought‐, high‐salt‐and cold‐responsive gene expression. The Plant Journal, vol. 33, no. 4, pp. 751-763. http://dx.doi.org/10.1046/j.1365-313X.2003.01661.x. PMid:12609047.
http://dx.doi.org/10.1046/j.1365-313X.20... ), OsDREB1F (Rehman & Mahmood, 2015REHMAN, S. and MAHMOOD, T., 2015. Functional role of DREB and ERF transcription factors: regulating stress-responsive network in plants. Acta Physiologiae Plantarum, vol. 37, no. 9, pp. 178. http://dx.doi.org/10.1007/s11738-015-1929-1.
http://dx.doi.org/10.1007/s11738-015-192... ), HvDRF1 (Xue and Loveridge, 2004XUE, G.P. and LOVERIDGE, C.W., 2004. HvDRF1 is involved in abscisic acid‐mediated gene regulation in barley and produces two forms of AP2 transcriptional activators, interacting preferably with a CT‐rich element. The Plant Journal, vol. 37, no. 3, pp. 326-339. http://dx.doi.org/10.1046/j.1365-313X.2003.01963.x. PMid:14731254.
http://dx.doi.org/10.1046/j.1365-313X.20... ), HvDREB1 (Xu et al., 2009XU, Z.S., NI, Z.Y., LI, Z.Y., LI, L.C., CHEN, M., GAO, D.Y. and MA, Y.Z., 2009. Isolation and functional characterization of HvDREB1: a gene encoding a dehydration-responsive element binding protein in Hordeum vulgare. Journal of Plant Research, vol. 122, no. 1, pp. 121-130. http://dx.doi.org/10.1007/s10265-008-0195-3. PMid:19067111.
http://dx.doi.org/10.1007/s10265-008-019... ), ZmDREB2A (Qin et al., 2007QIN, F., KAKIMOTO, M., SAKUMA, Y., MARUYAMA, K., OSAKABE, Y., TRAN, L.S.P. and YAMAGUCHI‐SHINOZAKI, K., 2007. Regulation and functional analysis of ZmDREB2A in response to drought and heat stresses in Zea mays L. The Plant Journal, vol. 50, no. 1, pp. 54-69. http://dx.doi.org/10.1111/j.1365-313X.2007.03034.x. PMid:17346263.
http://dx.doi.org/10.1111/j.1365-313X.20... ), SiDREB2 (Lata et al., 2011LATA, C., BHUTTY, S., BAHADUR, R.P., MAJEE, M. and PRASAD, M., 2011. Association of an SNP in a novel DREB2-like gene SiDREB2 with stress tolerance in foxtail millet [Setaria italica (L.)]. Journal of Experimental Botany, vol. 62, no. 10, pp. 3387-3401. http://dx.doi.org/10.1093/jxb/err016. PMid:21414959.
http://dx.doi.org/10.1093/jxb/err016... ),

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Figure 1
Abiotic stress Signal Transduction in Plant cell; Schematic representation of regulons (In plant cells) involves in abiotic stresses responses. Transcription factors and its stress inducible gene are showing in boxes.

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Figure 2
bZIP stress (drought and salt) inducible transcription factors in different plant. A family of bZIP induced the following genes under the influence of abiotic stress of drought and salt.ABF2 (Choi et al., 2000CHOI, H.I., HONG, J.H., HA, J.O., KANG, J.Y. and KIM, S.Y., 2000. ABFs, a family of ABA-responsive element binding factors. The Journal of Biological Chemistry, vol. 275, no. 3, pp. 1723-1730. http://dx.doi.org/10.1074/jbc.275.3.1723. PMid:10636868.
http://dx.doi.org/10.1074/jbc.275.3.1723... ), GmbZIP44, GmbZIP62 & GmbZIP78 (Liao et al., 2008cLIAO, Y., ZOU, H.F., WEI, W., HAO, Y.J., TIAN, A.G., HUANG, J. and CHEN, S.Y., 2008c. Soybean GmbZIP44, GmbZIP62 and GmbZIP78 genes function as negative regulator of ABA signaling and confer salt and freezing tolerance in transgenic Arabidopsis. Planta, vol. 228, no. 2, pp. 225-240. http://dx.doi.org/10.1007/s00425-008-0731-3. PMid:18365246.
http://dx.doi.org/10.1007/s00425-008-073... ), GmbZIP132 (Liao et al., 2008aLIAO, Y., ZHANG, J.S., CHEN, S.Y. and ZHANG, W.K., 2008a. Role of soybean GmbZIP132 under abscisic acid and salt stresses. Journal of Integrative Plant Biology, vol. 50, no. 2, pp. 221-230. http://dx.doi.org/10.1111/j.1744-7909.2007.00593.x. PMid:18713445.
http://dx.doi.org/10.1111/j.1744-7909.20... ), Wlip19 (Kobayashi et al., 2008KOBAYASHI, F., MAETA, E., TERASHIMA, A., KAWAURA, K., OGIHARA, Y. and TAKUMI, S., 2008. Development of abiotic stress tolerance via bZIP-type transcription factor LIP19 in common wheat. Journal of Experimental Botany, vol. 59, no. 4, pp. 891-905. http://dx.doi.org/10.1093/jxb/ern014. PMid:18326864.
http://dx.doi.org/10.1093/jxb/ern014... ), ZmbZIP17 (Jia et al., 2009JIA, Z., LIAN, Y., ZHU, Y., HE, J., CAO, Z. and WANG, G., 2009. Cloning and characterization of a putative transcription factor induced by abiotic stress in Zea mays. African Journal of Biotechnology, vol. 8, no. 24) and OsbZIP23 (Park et al. 2015PARK, S.H., JEONG, J.S., LEE, K.H., KIM, Y.S., DO CHOI, Y. and KIM, J.K., 2015. OsbZIP23 and OsbZIP45, members of the rice basic leucine zipper transcription factor family, are involved in drought tolerance. Plant Biotechnology Reports, vol. 9, no. 2, pp. 89-96. http://dx.doi.org/10.1007/s11816-015-0346-7.
http://dx.doi.org/10.1007/s11816-015-034... ).

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Figure 3
NAC gene family stress (drought and salt) inducible transcription factors in different plant. NAC regulon induced the following regulatory genes under the influence of abiotic stress of drought and salt. AtNAC019, AtNAC055 & AtNAC072 (Tran et al., 2004TRAN, L.S.P., NAKASHIMA, K., SAKUMA, Y., SIMPSON, S.D., FUJITA, Y., MARUYAMA, K. and YAMAGUCHI-SHINOZAKI, K., 2004. Isolation and functional analysis of Arabidopsis stress-inducible NAC transcription factors that bind to a drought-responsive cis-element in the early responsive to dehydration stress 1 promoter. The Plant Cell, vol. 16, no. 9, pp. 2481-2498. http://dx.doi.org/10.1105/tpc.104.022699. PMid:15319476.
http://dx.doi.org/10.1105/tpc.104.022699... ), SiNAC (Puranik et al., 2011PURANIK, S., BAHADUR, R.P., SRIVASTAVA, P.S. and PRASAD, M., 2011. Molecular cloning and characterization of a membrane associated NAC family gene, SiNAC from foxtail millet [Setaria italica (L.) P. Beauv.]. Molecular Biotechnology, vol. 49, no. 2, pp. 138-150. http://dx.doi.org/10.1007/s12033-011-9385-7. PMid:21312005.
http://dx.doi.org/10.1007/s12033-011-938... ), GhNAC4 (Meng et al., 2009MENG, C., CAI, C., ZHANG, T. and GUO, W., 2009. Characterization of six novel NAC genes and their responses to abiotic stresses in Gossypium hirsutum L. Plant Science, vol. 176, no. 3, pp. 352-359. http://dx.doi.org/10.1016/j.plantsci.2008.12.003.
http://dx.doi.org/10.1016/j.plantsci.200... ), SNAC1 (Hu et al., 2006HU, H., DAI, M., YAO, J., XIAO, B., LI, X., ZHANG, Q. and XIONG, L., 2006. Overexpressing a NAM, ATAF, and CUC (NAC) transcription factor enhances drought resistance and salt tolerance in rice. Proceedings of the National Academy of Sciences of the United States of America, vol. 103, no. 35, pp. 12987-12992. http://dx.doi.org/10.1073/pnas.0604882103. PMid:16924117.
http://dx.doi.org/10.1073/pnas.060488210... ), SNAC2 (Hu et al., 2008HU, H., YOU, J., FANG, Y., ZHU, X., QI, Z. and XIONG, L., 2008. Characterization of transcription factor gene SNAC2 conferring cold and salt tolerance in rice. Plant Molecular Biology, vol. 67, no. 1-2, pp. 169-181. http://dx.doi.org/10.1007/s11103-008-9309-5. PMid:18273684.
http://dx.doi.org/10.1007/s11103-008-930... ), OsNAC6 (Nakahsima et al., 2007; Rachmat et al., 2014RACHMAT, A., NUGROHO, S., SUKMA, D., ASWIDINNOOR, H. and SUDARSONO, S., 2014. Overexpression of OsNAC6 transcription factor from Indonesia rice cultivar enhances drought and salt tolerance. Emirates Journal of Food and Agriculture, vol. 26, no. 6, pp. 519-527. http://dx.doi.org/10.9755/ejfa.v26i6.17672.
http://dx.doi.org/10.9755/ejfa.v26i6.176... )

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Figure 4
MYB stress (drought and salt) inducible transcription factors in different plant. MYB stress (drought and salt) inducible transcription factors in different plant.OsMYB3R-2 (Dai et al., 2007DAI, X., XU, Y., MA, Q., XU, W., WANG, T., XUE, Y. and CHONG, K., 2007. Overexpression of an R1R2R3 MYB gene, OsMYB3R-2, increases tolerance to freezing, drought, and salt stress in transgenic Arabidopsis. Plant Physiology, vol. 143, no. 4, pp. 1739-1751. http://dx.doi.org/10.1104/pp.106.094532. PMid:17293435.
http://dx.doi.org/10.1104/pp.106.094532... ), AtMYB2 (Abe et al., 2003ABE, H., URAO, T., ITO, T., SEKI, M., SHINOZAKI, K. and YAMAGUCHI-SHINOZAKI, K., 2003. Arabidopsis AtMYC2 (bHLH) and AtMYB2 (MYB) function as transcriptional activators in abscisic acid signaling. The Plant Cell, vol. 15, no. 1, pp. 63-78. http://dx.doi.org/10.1105/tpc.006130. PMid:12509522.
http://dx.doi.org/10.1105/tpc.006130... ), GmMYB177 & GmMYB76 (Liao et al., 2008bLIAO, Y., ZOU, H.F., WANG, H.W., ZHANG, W.K., MA, B., ZHANG, J.S. and CHEN, S.Y., 2008b. Soybean GmMYB76, GmMYB92, and GmMYB177 genes confer stress tolerance in transgenic Arabidopsis plants. Cell Research, vol. 18, no. 10, pp. 1047-1060. http://dx.doi.org/10.1038/cr.2008.280. PMid:18725908.
http://dx.doi.org/10.1038/cr.2008.280... ) and MYB15 (Ding et al., 2009DING, Z., LI, S., AN, X., LIU, X., QIN, H. and WANG, D., 2009. Transgenic expression of MYB15 confers enhanced sensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana. Journal of Genetics and Genomics, vol. 36, no. 1, pp. 17-29. http://dx.doi.org/10.1016/S1673-8527(09)60003-5. PMid:19161942.
http://dx.doi.org/10.1016/S1673-8527(09)... )

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Table 1
Transgenic Arabidopsis plant using different stress (Salt and Drought) inducible Transcription factors.

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Table 2
Transgenic rice plant using different stress (Salt and Drought) inducible Transcription factors

Figure 5 DREB gene family express under drought and sat stress condition in different plants. GmDREBa, GmDREBb, GmDREBc (Li et al., 2005LI, X.P., TIAN, A.G., LUO, G.Z., GONG, Z.Z., ZHANG, J.S. and CHEN, S.Y., 2005. Soybean DRE-binding transcription factors that are responsive to abiotic stresses. Theoretical and Applied Genetics, vol. 110, no. 8, pp. 1355-1362. http://dx.doi.org/10.1007/s00122-004-1867-6. PMid:15841365.
http://dx.doi.org/10.1007/s00122-004-186... ), GmDREBa (Chen et al., 2007CHEN, M., WANG, Q.Y., CHENG, X.G., XU, Z.S., LI, L.C., YE, X.G., XIA, L.Q. and MA, Y.Z., 2007. GmDREB2, a soybean DRE-binding transcription factor, conferred drought and high-salt tolerance in transgenic plants. Biochemical and Biophysical Research Communications, vol. 353, no. 2, pp. 299-305. http://dx.doi.org/10.1016/j.bbrc.2006.12.027. PMid:17178106.
http://dx.doi.org/10.1016/j.bbrc.2006.12... ), CAP2 (Shukla et al., 2006SHUKLA, R.K., RAHA, S., TRIPATHI, V. and CHATTOPADHYAY, D., 2006. Expression of CAP2, an APETALA2-family transcription factor from chickpea, enhances growth and tolerance to dehydration and salt stress in transgenic tobacco. Plant Physiology, vol. 142, no. 1, pp. 113-123. http://dx.doi.org/10.1104/pp.106.081752. PMid:16844836.
http://dx.doi.org/10.1104/pp.106.081752... ), DREB2A (Liu et al., 1998LIU, Q., KASUGA, M., SAKUMA, Y., ABE, H., MIURA, S., YAMAGUCHI-SHINOZAKI, K. and SHINOZAKI, K., 1998. Two transcription factors, DREB1 and DREB2, with an EREBP/AP2 DNA binding domain separate two cellular signal transduction pathways in drought-and low-temperature-responsive gene expression, respectively, in Arabidopsis. The Plant Cell, vol. 10, no. 8, pp. 1391-1406. http://dx.doi.org/10.1105/tpc.10.8.1391. PMid:9707537.
http://dx.doi.org/10.1105/tpc.10.8.1391... ), OsDREB1C & OsDREB2A (Dubouzet et al., 2003DUBOUZET, J.G., SAKUMA, Y., ITO, Y., KASUGA, M., DUBOUZET, E.G., MIURA, S., SEKI, M., SHINOZAKI, K. and YAMAGUCHI-SHINOZAKI, K., 2003. OsDREB genes in rice, Oryza sativa L., encode transcription activators that function in drought‐, high‐salt‐and cold‐responsive gene expression. The Plant Journal, vol. 33, no. 4, pp. 751-763. http://dx.doi.org/10.1046/j.1365-313X.2003.01661.x. PMid:12609047.
http://dx.doi.org/10.1046/j.1365-313X.20... ), OsDREB1F (Rehman & Mahmood, 2015REHMAN, S. and MAHMOOD, T., 2015. Functional role of DREB and ERF transcription factors: regulating stress-responsive network in plants. Acta Physiologiae Plantarum, vol. 37, no. 9, pp. 178. http://dx.doi.org/10.1007/s11738-015-1929-1.
http://dx.doi.org/10.1007/s11738-015-192... ), HvDRF1 (Xue and Loveridge, 2004XUE, G.P. and LOVERIDGE, C.W., 2004. HvDRF1 is involved in abscisic acid‐mediated gene regulation in barley and produces two forms of AP2 transcriptional activators, interacting preferably with a CT‐rich element. The Plant Journal, vol. 37, no. 3, pp. 326-339. http://dx.doi.org/10.1046/j.1365-313X.2003.01963.x. PMid:14731254.
http://dx.doi.org/10.1046/j.1365-313X.20... ), HvDREB1 (Xu et al., 2009XU, Z.S., NI, Z.Y., LI, Z.Y., LI, L.C., CHEN, M., GAO, D.Y. and MA, Y.Z., 2009. Isolation and functional characterization of HvDREB1: a gene encoding a dehydration-responsive element binding protein in Hordeum vulgare. Journal of Plant Research, vol. 122, no. 1, pp. 121-130. http://dx.doi.org/10.1007/s10265-008-0195-3. PMid:19067111.
http://dx.doi.org/10.1007/s10265-008-019... ), ZmDREB2A (Qin et al., 2007QIN, F., KAKIMOTO, M., SAKUMA, Y., MARUYAMA, K., OSAKABE, Y., TRAN, L.S.P. and YAMAGUCHI‐SHINOZAKI, K., 2007. Regulation and functional analysis of ZmDREB2A in response to drought and heat stresses in Zea mays L. The Plant Journal, vol. 50, no. 1, pp. 54-69. http://dx.doi.org/10.1111/j.1365-313X.2007.03034.x. PMid:17346263.
http://dx.doi.org/10.1111/j.1365-313X.20... ), SiDREB2 (Lata et al., 2011LATA, C., BHUTTY, S., BAHADUR, R.P., MAJEE, M. and PRASAD, M., 2011. Association of an SNP in a novel DREB2-like gene SiDREB2 with stress tolerance in foxtail millet [Setaria italica (L.)]. Journal of Experimental Botany, vol. 62, no. 10, pp. 3387-3401. http://dx.doi.org/10.1093/jxb/err016. PMid:21414959.
http://dx.doi.org/10.1093/jxb/err016... ),

Figure 3 NAC gene family stress (drought and salt) inducible transcription factors in different plant. NAC regulon induced the following regulatory genes under the influence of abiotic stress of drought and salt. AtNAC019, AtNAC055 & AtNAC072 (Tran et al., 2004TRAN, L.S.P., NAKASHIMA, K., SAKUMA, Y., SIMPSON, S.D., FUJITA, Y., MARUYAMA, K. and YAMAGUCHI-SHINOZAKI, K., 2004. Isolation and functional analysis of Arabidopsis stress-inducible NAC transcription factors that bind to a drought-responsive cis-element in the early responsive to dehydration stress 1 promoter. The Plant Cell, vol. 16, no. 9, pp. 2481-2498. http://dx.doi.org/10.1105/tpc.104.022699. PMid:15319476.
http://dx.doi.org/10.1105/tpc.104.022699... ), SiNAC (Puranik et al., 2011PURANIK, S., BAHADUR, R.P., SRIVASTAVA, P.S. and PRASAD, M., 2011. Molecular cloning and characterization of a membrane associated NAC family gene, SiNAC from foxtail millet [Setaria italica (L.) P. Beauv.]. Molecular Biotechnology, vol. 49, no. 2, pp. 138-150. http://dx.doi.org/10.1007/s12033-011-9385-7. PMid:21312005.
http://dx.doi.org/10.1007/s12033-011-938... ), GhNAC4 (Meng et al., 2009MENG, C., CAI, C., ZHANG, T. and GUO, W., 2009. Characterization of six novel NAC genes and their responses to abiotic stresses in Gossypium hirsutum L. Plant Science, vol. 176, no. 3, pp. 352-359. http://dx.doi.org/10.1016/j.plantsci.2008.12.003.
http://dx.doi.org/10.1016/j.plantsci.200... ), SNAC1 (Hu et al., 2006HU, H., DAI, M., YAO, J., XIAO, B., LI, X., ZHANG, Q. and XIONG, L., 2006. Overexpressing a NAM, ATAF, and CUC (NAC) transcription factor enhances drought resistance and salt tolerance in rice. Proceedings of the National Academy of Sciences of the United States of America, vol. 103, no. 35, pp. 12987-12992. http://dx.doi.org/10.1073/pnas.0604882103. PMid:16924117.
http://dx.doi.org/10.1073/pnas.060488210... ), SNAC2 (Hu et al., 2008HU, H., YOU, J., FANG, Y., ZHU, X., QI, Z. and XIONG, L., 2008. Characterization of transcription factor gene SNAC2 conferring cold and salt tolerance in rice. Plant Molecular Biology, vol. 67, no. 1-2, pp. 169-181. http://dx.doi.org/10.1007/s11103-008-9309-5. PMid:18273684.
http://dx.doi.org/10.1007/s11103-008-930... ), OsNAC6 (Nakahsima et al., 2007; Rachmat et al., 2014RACHMAT, A., NUGROHO, S., SUKMA, D., ASWIDINNOOR, H. and SUDARSONO, S., 2014. Overexpression of OsNAC6 transcription factor from Indonesia rice cultivar enhances drought and salt tolerance. Emirates Journal of Food and Agriculture, vol. 26, no. 6, pp. 519-527. http://dx.doi.org/10.9755/ejfa.v26i6.17672.
http://dx.doi.org/10.9755/ejfa.v26i6.176... )

Transgenic plant	Transcription factor Family and gene in Transgenic approach		Stress tolerance	References
Arabidopsis	bZIP	ABF2	Drought	Kim et al. (2004)KIM, S., KANG, J.Y., CHO, D.I., PARK, J.H. and KIM, S.Y., 2004. ABF2, an ABRE‐binding bZIP factor, is an essential component of glucose signaling and its overexpression affects multiple stress tolerance. The Plant Journal, vol. 40, no. 1, pp. 75-87. http://dx.doi.org/10.1111/j.1365-313X.2004.02192.x. PMid:15361142. http://dx.doi.org/10.1111/j.1365-313X.20... ; Lee et al. (2010)LEE, S.J., KANG, J.Y., PARK, H.J., KIM, M.D., BAE, M.S., CHOI, H.I. and KIM, S.Y., 2010. DREB2C interacts with ABF2, a bZIP protein regulating abscisic acid-responsive gene expression, and its overexpression affects abscisic acid sensitivity. Plant Physiology, vol. 153, no. 2, pp. 716-727. http://dx.doi.org/10.1104/pp.110.154617. PMid:20395451. http://dx.doi.org/10.1104/pp.110.154617...
		ABF3	Drought	Kang et al. (2002KANG, J.Y., CHOI, H.I., IM, M.Y. and KIM, S.Y., 2002. Arabidopsis basic leucine zipper proteins that mediate stress-responsive abscisic acid signaling. The Plant Cell, vol. 14, no. 2, pp. 343-357. http://dx.doi.org/10.1105/tpc.010362. PMid:11884679. http://dx.doi.org/10.1105/tpc.010362... ; Yoshida et al. (2010)YOSHIDA, T., FUJITA, Y., SAYAMA, H., KIDOKORO, S., MARUYAMA, K., MIZOI, J., SHINOZAKI, K. and YAMAGUCHI-SHINOZAKI, K., 2010. AREB1, AREB2, and ABF3 are master transcription factors that cooperatively regulate ABRE‐dependent ABA signaling involved in drought stress tolerance and require ABA for full activation. The Plant Journal, vol. 61, no. 4, pp. 672-685. http://dx.doi.org/10.1111/j.1365-313X.2009.04092.x. PMid:19947981. http://dx.doi.org/10.1111/j.1365-313X.20...
		ABF4	Drought	Kang et al. (2002)KANG, J.Y., CHOI, H.I., IM, M.Y. and KIM, S.Y., 2002. Arabidopsis basic leucine zipper proteins that mediate stress-responsive abscisic acid signaling. The Plant Cell, vol. 14, no. 2, pp. 343-357. http://dx.doi.org/10.1105/tpc.010362. PMid:11884679. http://dx.doi.org/10.1105/tpc.010362... ; García et al. (2018)GARCÍA, M.N.M., CORTELEZZI, J.I., FUMAGALLI, M. and CAPIATI, D.A., 2018. Expression of the Arabidopsis ABF4 gene in potato increases tuber yield, improves tuber quality and enhances salt and drought tolerance. Plant Molecular Biology, vol. 98, no. 1-2, pp. 137-152. http://dx.doi.org/10.1007/s11103-018-0769-y. PMid:30143991. http://dx.doi.org/10.1007/s11103-018-076...
		GmbZIP44	Salinity	Liao et al., 2008c)LIAO, Y., ZOU, H.F., WEI, W., HAO, Y.J., TIAN, A.G., HUANG, J. and CHEN, S.Y., 2008c. Soybean GmbZIP44, GmbZIP62 and GmbZIP78 genes function as negative regulator of ABA signaling and confer salt and freezing tolerance in transgenic Arabidopsis. Planta, vol. 228, no. 2, pp. 225-240. http://dx.doi.org/10.1007/s00425-008-0731-3. PMid:18365246. http://dx.doi.org/10.1007/s00425-008-073... ; Cao et al. (2018)CAO, D., LI, Y., LIU, B., KONG, F. and TRAN, L.S.P., 2018. Adaptive mechanisms of soybean grown on salt‐affected soils. Land Degradation & Development, vol. 29, no. 4, pp. 1054-1064. http://dx.doi.org/10.1002/ldr.2754. http://dx.doi.org/10.1002/ldr.2754...
		GmbZIP62	Salinity	Liao et al., 2008cLIAO, Y., ZOU, H.F., WEI, W., HAO, Y.J., TIAN, A.G., HUANG, J. and CHEN, S.Y., 2008c. Soybean GmbZIP44, GmbZIP62 and GmbZIP78 genes function as negative regulator of ABA signaling and confer salt and freezing tolerance in transgenic Arabidopsis. Planta, vol. 228, no. 2, pp. 225-240. http://dx.doi.org/10.1007/s00425-008-0731-3. PMid:18365246. http://dx.doi.org/10.1007/s00425-008-073... ; Alves et al. (2015)ALVES, M.S., SOARES, Z.G., VIDIGAL, P.M., BARROS, E.G., PODDANOSQUI, A.M., AOYAGI, L.N., ABDELNOOR, R.V., MARCELINO-GUIMARÃES, F.C. and FIETTO, L.G., 2015. Differential expression of four soybean bZIP genes during Phakopsora pachyrhizi infection. Functional & Integrative Genomics, vol. 15, no. 6, pp. 685-696. http://dx.doi.org/10.1007/s10142-015-0445-0. PMid:26013145. http://dx.doi.org/10.1007/s10142-015-044...
		GmbZIP78	Salinity	Liao et al., 2008c)LIAO, Y., ZOU, H.F., WEI, W., HAO, Y.J., TIAN, A.G., HUANG, J. and CHEN, S.Y., 2008c. Soybean GmbZIP44, GmbZIP62 and GmbZIP78 genes function as negative regulator of ABA signaling and confer salt and freezing tolerance in transgenic Arabidopsis. Planta, vol. 228, no. 2, pp. 225-240. http://dx.doi.org/10.1007/s00425-008-0731-3. PMid:18365246. http://dx.doi.org/10.1007/s00425-008-073... ; Wang et al. (2018)WANG, L., ZHU, J., LI, X., WANG, S. and WU, J., 2018. Salt and drought stress and ABA responses related to bZIP genes from V. radiata and V. angularis. Gene, vol. 651, pp. 152-160. http://dx.doi.org/10.1016/j.gene.2018.02.005. PMid:29425824. http://dx.doi.org/10.1016/j.gene.2018.02...
		GmbZIP132	Salinity	Liao et al., 2008a)LIAO, Y., ZHANG, J.S., CHEN, S.Y. and ZHANG, W.K., 2008a. Role of soybean GmbZIP132 under abscisic acid and salt stresses. Journal of Integrative Plant Biology, vol. 50, no. 2, pp. 221-230. http://dx.doi.org/10.1111/j.1744-7909.2007.00593.x. PMid:18713445. http://dx.doi.org/10.1111/j.1744-7909.20... ; Hossain et al. (2015)HOSSAIN, M. R., VICKERS, L., SHARMA, G., LIVERMORE, T., PRITCHARD, J., and FORD-LLOYD, B. V., 2015. Salinity tolerance in plants. In: S.H. WANI and M.A. HOSSAIN. Managing salt tolerance in plants: molecular and genomic perspectives. Boca Raton: CRC Press, pp. 407-426.
		AtbZIP60	Salinity	Fujita et al. (2007)FUJITA, M., MIZUKADO, S., FUJITA, Y., ICHIKAWA, T., NAKAZAWA, M., SEKI, M. and SHINOZAKI, K., 2007. Identification of stress-tolerance-related transcription-factor genes via mini-scale Full-length cDNA Over-eXpressor (FOX) gene hunting system. Biochemical and Biophysical Research Communications, vol. 364, no. 2, pp. 250-257. http://dx.doi.org/10.1016/j.bbrc.2007.09.124. PMid:17937930. http://dx.doi.org/10.1016/j.bbrc.2007.09... ; Tang et al. (2012)TANG, W., FEI, Y. and PAGE, M., 2012. Elevated tolerance to salt stress in transgenic cells expressing transcription factor AtbZIP60 is associated with the increased activities of H+-ATPase and acid phosphatase. Plant Biotechnology Reports, vol. 6, no. 4, pp. 313-325. http://dx.doi.org/10.1007/s11816-012-0226-3. http://dx.doi.org/10.1007/s11816-012-022...
	MYB	MYB15	Drought, Salinity	Ding et al. (2009)DING, Z., LI, S., AN, X., LIU, X., QIN, H. and WANG, D., 2009. Transgenic expression of MYB15 confers enhanced sensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana. Journal of Genetics and Genomics, vol. 36, no. 1, pp. 17-29. http://dx.doi.org/10.1016/S1673-8527(09)60003-5. PMid:19161942. http://dx.doi.org/10.1016/S1673-8527(09)... ; Hou et al. (2018)HOU, H., JIA, H., YAN, Q. and WANG, X., 2018. Overexpression of a SBP-box gene (VpSBP16) from chinese wild vitis species in Arabidopsis improves salinity and drought stress tolerance. International Journal of Molecular Sciences, vol. 19, no. 4, pp. 940. http://dx.doi.org/10.3390/ijms19040940. PMid:29565279. http://dx.doi.org/10.3390/ijms19040940...
		OsMYB3R-2	Drought, Salinity	Dai et al. (2007)DAI, X., XU, Y., MA, Q., XU, W., WANG, T., XUE, Y. and CHONG, K., 2007. Overexpression of an R1R2R3 MYB gene, OsMYB3R-2, increases tolerance to freezing, drought, and salt stress in transgenic Arabidopsis. Plant Physiology, vol. 143, no. 4, pp. 1739-1751. http://dx.doi.org/10.1104/pp.106.094532. PMid:17293435. http://dx.doi.org/10.1104/pp.106.094532... ; Yang et al. (2012)YANG, A., DAI, X. and ZHANG, W.H., 2012. A R2R3-type MYB gene, OsMYB2, is involved in salt, cold, and dehydration tolerance in rice. Journal of Experimental Botany, vol. 63, no. 7, pp. 2541-2556. http://dx.doi.org/10.1093/jxb/err431. PMid:22301384. http://dx.doi.org/10.1093/jxb/err431...
		OsMYB4	Drought	Mattana et al. (2005)MATTANA, M., BIAZZI, E., CONSONNI, R., LOCATELLI, F., VANNINI, C., PROVERA, S. and CORAGGIO, I., 2005. Overexpression of Osmyb4 enhances compatible solute accumulation and increases stress tolerance of Arabidopsis thaliana. Physiologia Plantarum, vol. 125, no. 2, pp. 212-223. http://dx.doi.org/10.1111/j.1399-3054.2005.00551.x. http://dx.doi.org/10.1111/j.1399-3054.20... ; Raldugina et al. (2018)RALDUGINA, G.N., MAREE, M., MATTANA, M., SHUMKOVA, G., MAPELLI, S., KHOLODOVA, V.P. and KUZNETSOV, V.V., 2018. Expression of rice OsMyb4 transcription factor improves tolerance to copper or zinc in canola plants. Biologia Plantarum, vol. 62, no. 3, pp. 511-520. http://dx.doi.org/10.1007/s10535-018-0800-9. http://dx.doi.org/10.1007/s10535-018-080...
	CBF/DREB	AtDREB1A	Drought	Liu et al. (1998)LIU, Q., KASUGA, M., SAKUMA, Y., ABE, H., MIURA, S., YAMAGUCHI-SHINOZAKI, K. and SHINOZAKI, K., 1998. Two transcription factors, DREB1 and DREB2, with an EREBP/AP2 DNA binding domain separate two cellular signal transduction pathways in drought-and low-temperature-responsive gene expression, respectively, in Arabidopsis. The Plant Cell, vol. 10, no. 8, pp. 1391-1406. http://dx.doi.org/10.1105/tpc.10.8.1391. PMid:9707537. http://dx.doi.org/10.1105/tpc.10.8.1391... ; Oh et al. (2005)OH, S.J., SONG, S.I., KIM, Y.S., JANG, H.J., KIM, S.Y., KIM, M., KIM, J.K., NAHM, B.H. and KIM, J.K., 2005. Arabidopsis CBF3/DREB1A and ABF3 in transgenic rice increased tolerance to abiotic stress without stunting growth. Plant Physiology, vol. 138, no. 1, pp. 341-351. http://dx.doi.org/10.1104/pp.104.059147. PMid:15834008. http://dx.doi.org/10.1104/pp.104.059147...
		AtDREB1A	Drought	Rehman & Mahmood (2015)REHMAN, S. and MAHMOOD, T., 2015. Functional role of DREB and ERF transcription factors: regulating stress-responsive network in plants. Acta Physiologiae Plantarum, vol. 37, no. 9, pp. 178. http://dx.doi.org/10.1007/s11738-015-1929-1. http://dx.doi.org/10.1007/s11738-015-192...
		AtDREB2A	Drought	Rehman & Mahmood (2015)
		AtCBF1	Salinity	Ma et al. (2015)MA, X., ZHU, X., LI, C., SONG, Y., ZHANG, W., XIA, G. and WANG, M., 2015. Overexpression of wheat NF-YA10 gene regulates the salinity stress response in Arabidopsis thaliana. Plant Physiology and Biochemistry, vol. 86, pp. 34-43. http://dx.doi.org/10.1016/j.plaphy.2014.11.011. PMid:25461698. http://dx.doi.org/10.1016/j.plaphy.2014....
		OsDREB1A	Drought, Salinity,	Dubouzet et al. (2003)DUBOUZET, J.G., SAKUMA, Y., ITO, Y., KASUGA, M., DUBOUZET, E.G., MIURA, S., SEKI, M., SHINOZAKI, K. and YAMAGUCHI-SHINOZAKI, K., 2003. OsDREB genes in rice, Oryza sativa L., encode transcription activators that function in drought‐, high‐salt‐and cold‐responsive gene expression. The Plant Journal, vol. 33, no. 4, pp. 751-763. http://dx.doi.org/10.1046/j.1365-313X.2003.01661.x. PMid:12609047. http://dx.doi.org/10.1046/j.1365-313X.20...
		OsDREB2B	Drought,	Matsukura et al. (2010)MATSUKURA, S., MIZOI, J., YOSHIDA, T., TODAKA, D., ITO, Y., MARUYAMA, K., SHINOZAKI, K. and YAMAGUCHI-SHINOZAKI, K., 2010. Comprehensive analysis of rice DREB2-type genes that encode transcription factors involved in the expression of abiotic stress-responsive genes. Molecular Genetics and Genomics, vol. 283, no. 2, pp. 185-196. http://dx.doi.org/10.1007/s00438-009-0506-y. PMid:20049613. http://dx.doi.org/10.1007/s00438-009-050... ; Joshi et al. (2016)JOSHI, R., WANI, S.H., SINGH, B., BOHRA, A., DAR, Z.A., LONE, A.A. and SINGLA-PAREEK, S.L., 2016. Transcription factors and plants response to drought stress: current understanding and future directions. Frontiers of Plant Science, vol. 7, pp. 1029. http://dx.doi.org/10.3389/fpls.2016.01029. PMid:27471513. http://dx.doi.org/10.3389/fpls.2016.0102...
		OsDREB1F	Drought, Salinity,	Rehman & Mahmood (2015)
		ZmDREB2A	Drought	Qin et al. (2007)QIN, F., KAKIMOTO, M., SAKUMA, Y., MARUYAMA, K., OSAKABE, Y., TRAN, L.S.P. and YAMAGUCHI‐SHINOZAKI, K., 2007. Regulation and functional analysis of ZmDREB2A in response to drought and heat stresses in Zea mays L. The Plant Journal, vol. 50, no. 1, pp. 54-69. http://dx.doi.org/10.1111/j.1365-313X.2007.03034.x. PMid:17346263. http://dx.doi.org/10.1111/j.1365-313X.20...
		HvDREB1	Salinity	Xu et al. (2009)XU, Z.S., NI, Z.Y., LI, Z.Y., LI, L.C., CHEN, M., GAO, D.Y. and MA, Y.Z., 2009. Isolation and functional characterization of HvDREB1: a gene encoding a dehydration-responsive element binding protein in Hordeum vulgare. Journal of Plant Research, vol. 122, no. 1, pp. 121-130. http://dx.doi.org/10.1007/s10265-008-0195-3. PMid:19067111. http://dx.doi.org/10.1007/s10265-008-019...
	NAC	AtNAC2	Drought	Tran et al. (2004)TRAN, L.S.P., NAKASHIMA, K., SAKUMA, Y., SIMPSON, S.D., FUJITA, Y., MARUYAMA, K. and YAMAGUCHI-SHINOZAKI, K., 2004. Isolation and functional analysis of Arabidopsis stress-inducible NAC transcription factors that bind to a drought-responsive cis-element in the early responsive to dehydration stress 1 promoter. The Plant Cell, vol. 16, no. 9, pp. 2481-2498. http://dx.doi.org/10.1105/tpc.104.022699. PMid:15319476. http://dx.doi.org/10.1105/tpc.104.022699...
		AtNAC019	Drought	Tran et al. (2004)TRAN, L.S.P., NAKASHIMA, K., SAKUMA, Y., SIMPSON, S.D., FUJITA, Y., MARUYAMA, K. and YAMAGUCHI-SHINOZAKI, K., 2004. Isolation and functional analysis of Arabidopsis stress-inducible NAC transcription factors that bind to a drought-responsive cis-element in the early responsive to dehydration stress 1 promoter. The Plant Cell, vol. 16, no. 9, pp. 2481-2498. http://dx.doi.org/10.1105/tpc.104.022699. PMid:15319476. http://dx.doi.org/10.1105/tpc.104.022699...
		AtNAC055	Drought	Tran et al. (2004)TRAN, L.S.P., NAKASHIMA, K., SAKUMA, Y., SIMPSON, S.D., FUJITA, Y., MARUYAMA, K. and YAMAGUCHI-SHINOZAKI, K., 2004. Isolation and functional analysis of Arabidopsis stress-inducible NAC transcription factors that bind to a drought-responsive cis-element in the early responsive to dehydration stress 1 promoter. The Plant Cell, vol. 16, no. 9, pp. 2481-2498. http://dx.doi.org/10.1105/tpc.104.022699. PMid:15319476. http://dx.doi.org/10.1105/tpc.104.022699...
	WRKY	OsWRKY45	Drought, Salinity	Qiu and Yu (2009)QIU, Y. and YU, D., 2009. Over-expression of the stress-induced OsWRKY45 enhances disease resistance and drought tolerance in Arabidopsis. Environmental and Experimental Botany, vol. 65, no. 1, pp. 35-47. http://dx.doi.org/10.1016/j.envexpbot.2008.07.002. http://dx.doi.org/10.1016/j.envexpbot.20...
	WRKY	GmWRKY54	Drought, Salinity	Tripathi et al. (2014)TRIPATHI, P., RABARA, R.C. and RUSHTON, P.J., 2014. A systems biology perspective on the role of WRKY transcription factors in drought responses in plants. Planta, vol. 239, no. 2, pp. 255-266. http://dx.doi.org/10.1007/s00425-013-1985-y. PMid:24146023. http://dx.doi.org/10.1007/s00425-013-198...

Transgenic plant	Transcription factor Family and gene in Transgenic approach		Stress tolerance	References
Rice	bZIP	OsABI5	Salinity	Zou et al. (2008)ZOU, M., GUAN, Y., REN, H., ZHANG, F. and CHEN, F., 2008. A bZIP transcription factor, OsABI5, is involved in rice fertility and stress tolerance. Plant Molecular Biology, vol. 66, no. 6, pp. 675-683. http://dx.doi.org/10.1007/s11103-008-9298-4. PMid:18236009. http://dx.doi.org/10.1007/s11103-008-929...
		OsbZIP23	Drought, Salinity	Xiang et al. (2008)XIANG, Y., TANG, N., DU, H., YE, H. and XIONG, L., 2008. Characterization of OsbZIP23 as a key player of the basic leucine zipper transcription factor family for conferring abscisic acid sensitivity and salinity and drought tolerance in rice. Plant Physiology, vol. 148, no. 4, pp. 1938-1952. http://dx.doi.org/10.1104/pp.108.128199. PMid:18931143. http://dx.doi.org/10.1104/pp.108.128199...
		OsAREB1	Drought	Jin et al. (2010)JIN, X.F., XIONG, A.S., PENG, R.H., LIU, J.G., GAO, F., CHEN, J.M. and YAO, Q.H., 2010. OsAREB1, an ABRE-binding protein responding to ABA and glucose, has multiple functions in Arabidopsis. BMB Reports, vol. 43, no. 1, pp. 34-39. http://dx.doi.org/10.5483/BMBRep.2010.43.1.034. PMid:20132733. http://dx.doi.org/10.5483/BMBRep.2010.43...
	CBF/DREB	AtDREB1A	Drought, Salinity	Oh et al. (2005)OH, S.J., SONG, S.I., KIM, Y.S., JANG, H.J., KIM, S.Y., KIM, M., KIM, J.K., NAHM, B.H. and KIM, J.K., 2005. Arabidopsis CBF3/DREB1A and ABF3 in transgenic rice increased tolerance to abiotic stress without stunting growth. Plant Physiology, vol. 138, no. 1, pp. 341-351. http://dx.doi.org/10.1104/pp.104.059147. PMid:15834008. http://dx.doi.org/10.1104/pp.104.059147...
		OsDREB1F	Drought, Salinity,	Rehman & Mahmood (2015)REHMAN, S. and MAHMOOD, T., 2015. Functional role of DREB and ERF transcription factors: regulating stress-responsive network in plants. Acta Physiologiae Plantarum, vol. 37, no. 9, pp. 178. http://dx.doi.org/10.1007/s11738-015-1929-1. http://dx.doi.org/10.1007/s11738-015-192...
		OsDREB1G	Drought	Chen et al. (2008)CHEN, J.Q., MENG, X.P., ZHANG, Y., XIA, M. and WANG, X.P., 2008. Over-expression of OsDREB genes lead to enhanced drought tolerance in rice. Biotechnology Letters, vol. 30, no. 12, pp. 2191-2198. http://dx.doi.org/10.1007/s10529-008-9811-5. PMid:18779926. http://dx.doi.org/10.1007/s10529-008-981...
	NAC	SNAC1	Drought, Salinity	Hu et al. (2006)HU, H., DAI, M., YAO, J., XIAO, B., LI, X., ZHANG, Q. and XIONG, L., 2006. Overexpressing a NAM, ATAF, and CUC (NAC) transcription factor enhances drought resistance and salt tolerance in rice. Proceedings of the National Academy of Sciences of the United States of America, vol. 103, no. 35, pp. 12987-12992. http://dx.doi.org/10.1073/pnas.0604882103. PMid:16924117. http://dx.doi.org/10.1073/pnas.060488210...
		OsNAC6	Drought, Salinity	Nakashima et al. (2007) NAKASHIMA, K., TRAN, L. S. P., VAN NGUYEN, D., FUJITA, M., MARUYAMA, K., TODAKA, D., and YAMAGUCHI‐SHINOZAKI, K., 2007. Functional analysis of a NAC‐type transcription factor OsNAC6 involved in abiotic and biotic stress‐responsive gene expression in rice.The Plant Journal, vol. 51, no. 4, pp. 617-630.
		ONAC045	Drought, Salinity	Zheng et al. (2009)ZHENG, X., CHEN, B., LU, G. and HAN, B., 2009. Overexpression of a NAC transcription factor enhances rice drought and salt tolerance.Biochemical and Biophysical Research Communications, 379(4), pp. 985-989.
	Others	HARDY	Drought, Salinity	Karaba et al. (2007)KARABA, A., DIXIT, S., GRECO, R., AHARONI, A., TRIJATMIKO, K.R., MARSCH-MARTINEZ, N. and PEREIRA, A., 2007. Improvement of water use efficiency in rice by expression of HARDY, an Arabidopsis drought and salt tolerance gene. Proceedings of the National Academy of Sciences of the United States of America, vol. 104, no. 39, pp. 15270-15275. http://dx.doi.org/10.1073/pnas.0707294104. PMid:17881564. http://dx.doi.org/10.1073/pnas.070729410... ; Guo et al. (2017)GUO, X., ZHANG, L., ZHU, J., WANG, A. and LIU, H., 2017. Christolea crassifolia HARDY gene enhances drought stress tolerance in transgenic tomato plants. Plant Cell, Tissue and Organ Culture, vol. 129, no. 3, pp. 469-481. http://dx.doi.org/10.1007/s11240-017-1192-9. http://dx.doi.org/10.1007/s11240-017-119...

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Regulon: Uma visão geral do sistema regulador da transcrição do estresse abiótico em plantas e papel nas plantas transgênica

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