Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. rwt FI Drn,Tetr ite,Lc,Ce oe,20,Ktk ot agl,Bce von & Bicker Ganguli, tran- Port, for Kotak, needed 2000, is Nover, which & region, Chen protein terminal Lyck, binding TATA pro- C Kistner, complex, export Treuter, one the transcription (Doring, and at possess basal TFIIB the HSFs NES import with with A to nuclear interacting or the Class close regulate by and located HSFs probably 2001). (NLS) activation, (AHA) Nover, A scriptional signal motifs & class activator in localization Winkelhaus Gagliardi, of Scharf, more nuclear Bonzelius, either C-terminus (Nover, or Doring, The localized ODs the (Heerklotz, their proteins, at conditions. respectively of shuttling (NES) cellular cesses, features are the signal the factors on on export shock based depending nuclear is Heat nuclei, classes or C 1996). cytoplasm and Gurley, and B Classifica- DBD & A, to Czarnecka-Verner activation. next into transcriptional Vergne, located HSFs during is plant trimerisation motif) of and HR-A/B interactions tion or protein-protein (OD Scharf for Domain 2010, responsible Oligomerisation Enoki, The is 5’-nGAAnnTTCn- & 1996). (HSE, element (Sakurai Ruterjans, shock & promoters heat lo- Nover the target is recognize (DBD) 2010, the which genome. domain protein, of Enoki, plant binding the DNA a 3’) & of Their in N-terminus Sakurai HSFs the domains. conserved to 2016, 25.7 well close of various Lu, cated average with & an structure modular with Gong a species, have Luo, plant HSFs HSFs, (http://www.cibiv.at/services/hsf/) Ma, 33 database encode from Liu, members Heatster HSF sequences 52 (Guo, The HSF single to 848 classes 2012). a 18 contains Nover, C currently with have & and families Drosophila Ebersberger B gene and Berberich, large A, Scharf, plants yeast into in While divided shock HSFs, are four heat 1996). which with encoding maintain live Vierling, and genes cells & temperatures mammalian of high Viitanen and during regulators damage (Boston, transcriptional from homeostasis cells as protect protein identified that chaperons been molecular have (HSPs), proteins (HSFs) adverse factors multiple shock to Heat adaptation the facilitates INTRODUCTION HSFs plant 1. in change. Diversity climate to HSF. response plant which complexity in the a proteins feature illustrate important with to death, an HSFA4A regulators, related cell conditions, Arabidopsis programmed transcription networks environmental the and including regulatory stresses of proteins different the Interactions homomeric regulatory to of of responses other (HSPs). homeostasis, formation provides of proteins cellular include number modifications transcription, shock which a control posttranslational heat interactions with and or protein-protein complexes proteins of splicing differences and web chromatin-associated considerable alternative trimers, complex with Moreover, HSF in control heteromeric involved expression genes. and are in HSF the HSFs reflected of individual toxic variability. is salinity, overview of further soil Variability an regulate conditions, profiles can provide hypoxic defenses. drought, HSFs we transcript to pathogen temperatures, interactions. Here only low in or and or not regulation threats. irradiation high versatility, responses including pathogen strong functional of conditions and their minerals, extreme regulators of of stresses primary number analysis abiotic are a through other to and HSFs tolerance of plant families number the gene of a large world to by diverse also encoded but are temperatures (HSFs) Factors high Shock Heat plants In Abstract 2020 20, July 1 Andr´asiNorbert and interactions regulation, factors: functions. shock heat plant of Variability ilgclRsac Centre Research Biological 1 Alad´ar Pettk´o-Szandtner, 1 n azoSzabados Laszlo and , 1 tal. et 02 cutes,Knr,Gs,Scharf, Gase, Kunert, Schultheiss, 2012, , 1 Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. i,Yn,L hn,21)o ev eas(a,Zag u i i hme,Xa h,Zo,Reiter, heat Zhou, and Shi, drought Xia, enhanced Ahammed, and Li, signaling Qi, peroxide 2018), Xu, promoted Zhang, Shen, overexpression (Cai, HSFA1B & metals Yang heavy 2017). Chen, Yan, or (Qian, Zhou, Zhou, 2014) treatments & oxidative Zhang, Liu, Yu 2013), & Wang, Beynon, Mullineaux, Li Mishra Martin, Zhu, Yang, & Hickman, 1995, 2020, Ott Liu, Bowden, Schoffl, Schoffl, Persad, Jan, Morison, Richard, & & Sparrow, Baker, Fryer, Buchanan-Wollaston, Hubel Ahmed Tanaka, Lawson, Lee, Iqbal, Albihlal, Ishida, includ- (Bechtold, 2013, Khan, Yamaguchi-Shinozaki, drought stresses Taji, Munir, Kusakabe, several & Ali, Shimura, to Hayashi (Liu Shah, tolerance Katori, Sakata, also conferred Ishikawa, stresses Shinozaki, species Ohama, other Seki, different (Higashi, to from heat responses HSFA1 Reduced control ing of but overexpression defects. tolerance, hand, developmental H heat other and and regulate salt growth many only retarded and not heat displayed (QK, most mutants and of knockout thermotolerance, expression HSFA1 impaired Arabidopsis cytoplasmic showed Quadruple misfolded to ) analysis. attach Scharf genetic to 2002, removed their are Scharf, encumbers HSPs & as conditions Nover (Scharf stress Theres, proteins during Tschiersch, released are Winkelhaus, and Tripp, HSP70/90 Mishra, 2011, (Driedonks al. S1) Charng, et (Table & genes Liao heat-induced Liu, revealed of activation which transcription reports early on made stresses HSFA1-type was abiotic to Focus responses Guo of 2015, S1). Regulation responses. 2.1 Rieu, (Table stress & heat factors from Park investigated different Peters, functions the Xu, responses biological of control (Driedonks, can functions S1) but (Table biological tolerance constraints Scharf heat abiotic 2016, regulate and only , not biotic HSFs many plant to PLANTS that IN demonstrate FACTORS reports SHOCK Numerous HEAT OF interactions. DIVERSITY protein-protein and FUNCTIONAL modifications 2. transcriptional post (Guo expression, Wang reviews in 2007, plant variation excellent of Nover, summarizing evolution in & and described Scharf features Structural been Koskull-Doring, partners. HSFs environments. thoroughly protein land of have of variable divergence to HSFs most spectrum functional adaptation in the wide with facilitated variable are and with associated memers are DBDs interact is ODs domains domains, can (Wang activation evolution while protein HSFs during and the conservation, individual oligomerisation Among that functional of few suggesting Diversification for for duplications 2018). sequence, pressure genome exceptions Xu, lethal and whole evolutionary with & size and not strong Ma HSFs, gene are through implying Chen, other mutants happened Shi, conserved, by Hsf HSFs (Wang, complemented plant evolution of be diversification. during Multiplication cannot functional genes. which considerable related phenotypes closely with protein particular 2010). of families have Enoki, maintenance & HSF include but (Sakurai functions large conditions HSF physiological have of standard range under Plants Broad also but metabolism, 2015). stress Dinkova-Kostova, Lee, of during & & only Naidu control (Dayalan Cha not to proteins, carcinogenesis controls Gil, homeostasis cellular damaged cells Lee, lead and mammalian Kim, of stresses or Kang, cellular in elimination of 2017, lethal HSF1 repair, range to broad are 1988). DNA a Pelham, unrelated mutations in to & responses mechanisms functioning HSF Sorger genes cellular 1997, Drosophila Wu, of control or & transcription In they Yeast Mortin (Jedlicka, that known. responses functions. yet suggesting stress species. multiple not abnormalities, those have is developmental in HSFs’ to functions considerable C specialized shown the possess Class were protein, to of HSFs the suggesting function function expanded of Precise repressor is region transcriptional family regulatory 2004). their this the Gurley, with monocots in & associated domain is Salem which repressor Pan, motif, (Czarnecka-Verner, a LFGV have tetrapeptide HSFs conserved B highly Class 2004). Koskull-Doring, 02.I o-tescniin SA atr r ehrdt nciectpamccmlxswith complexes cytoplasmic inactive to tethered are factors HSFA1 conditions non-stress In 2012). , rncito atr Ts r osdrda atrrgltro hrooeac,directing thermotolerance, of regulator master as considered are (TFs) factors transcription tal. et tal. et 02 aua 06.W aecmie ulctoso ln Sswihreport which HSFs plant on publications compiled have We 2016). Yabuta, 2012, , 02.Pat aesvrlHF1gnswt atal vrapn ucin,which functions, overlapping partially with genes HSFA1 several have Plants 2012). , tal. et 08.Poieaino h S eefml edt pcaiaino individual of specialization to lead family gene HSF the of Proliferation 2018). , 2 O tal. et 2 idcdgnsi Kmtn niae htHF1genes HSFA1 that indicated mutant QK in genes -induced 2 08.Hr efcso hi ucinldiversity, functional their on focus we Here 2018). , tal. et 06 Scharf 2016, , tal. et 05 Guo 2015, , tal. et tal. et tal. et athsfa1a,1b,1d,1e 01.O the On 2011). , 02 Shah, 2002, , tal. et 02 von 2012, , 2016, , tal. et Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. tesmmr eiso pgntcrglto n novshsoemtyaino agtpooes(Lamke, promoters target associated of transiently (Charng methylation was histone memory in involves it stress factor and as regulation stress heat Yang, epigenetic versatile thermotolerance with Huang, on and extend relies Wang, associated important to memory 2019, genes an stress described Sheteiwy, is of HSFA2 was & promoters HSFA2 that Guo with demonstrate Zhao, results species. Zhang, These plant heat Zhang, the different 2016). enhanced Wang, complement Huang, & bermudagrass Shao, could & Xu African and Zhang, Chai, Liu from plants, Zhou, Li, CtHsfA2b Arabidopsis Chen, 2018, and transgenic He, (Jiang, Yu, maize of (Li, of tolerance conditions ZmHSFA05 stress- salt germination osmotic and improved and induce high & which ZmHSF04 in and soybean Iwabuchi of Arabidopsis 2014). Hirochika, tolerance HSF transgenic Ma, A2-type Matsui, salinity in drought-induced growth and enhance Kondou, root heat Ichikawa, could a and (Yokotani, is rice GmHSF-34 Arabidopsis from 2008). transgenic Oda, OsHSFA2e or in as Schramm GolS1), genes species, 2009, 1, associated Shigeoka, synthase among & galactinol conserved Yabuta scavengers is Yoshida, (eg. ROS Nishizawa-Yokoi, enzymes Hsp18.1-CI), BAG6)(Fragkostefanakis 2006, metabolic (eg. 6, , protective athanogene HSPs APX2), (Bcl-2-associated included regulators which 2, apoptotic genes, peroxidase target ascorbate 2013). of Charng, spectrum (eg. & broad (Liu a cascade controls regulatory HSFA2 the in HSFA1 of downstream is H and heat defects, Pucciariello (Nishizawa viologen the methyl hand and 2012), other light al. Perata, high the also, & by On Banti generated heteroligomers Pucciariello, 2006). stresses anoxia oxidative 2010, form , submergence, and Perata, 2007), they heat & simultaneous Nishiuchi, Alpi but to & Loreti, Yamaguchi and Mafessoni, Koskull- factors, (Ogawa, (Banti, stresses HSFA1 von conditions osmotic oxidative by & and and Walch salt induced to Englich, is tolerance Kiehlmann, enhance HSFA2 (Scharf Bublak, Ganguli, of complexes Paul, Heider, superactivating Schramm, Expression Scharf, Simm, generating 1998, 2006, Fragkostefanakis, Shigeoka, Nover, 2006). 2007, & & Wang, Yoshimura Doring, Maruta, Schmidt & Yoshida, Chang Lyck, Yabuta, Nishizawa, Wang, Hohfeld, 2015, Chi, Schleiff, Liu, & Liu, Scharf (Charng, S1) (Table other 2016, Kusakabe, and Yamaguchi-Shinozaki, (Ohama, HSFA2 heat 2017). & conditions Yamaguchi-Shinozaki, transmit Shinozaki adverse & Yanagisawa, which in Shinozaki Ishida, cascade, homeostasis Sato, Takahashi, cellular transcriptional Ohama, Koizumi, maintain a Kidokoro, to overexpressing in Zhao, responsible in HSFA1 components Mizoi, is temperatures decisive 2015). and high (Li, and signals damage and first stress oxidative drought diminishing the and to are capacity tolerance by factors antioxidant levels enhancing increased (Cai by melatonin maize plants (COMT1) increasing of 1 Arabidopsis and O-methyltransferase ZmHSF06 acid genes A1-type caffeic HSP (Mishra conditions The gene tolerance activating genes biosynthetic light promotes through target melatonin HSFA1 high of tolerance the Tomato activation in stimulating cadmium proper 2013). APX2 confer for Chory, HSFA2 activated can requires & HSFA3 SlHSfA1a which Pogson and stresses, Functional Mockler, HSFA2 abiotic the several Hong, with defines HSFs 3). to Cole, which A together (Figure Class Estavillo, (DBD), HSFA1D, domain Crisp, tolerance 2019). Fragkostefanakis, binding alone. (Jung, stress & DNA Schleiff act in Scharf, of not Simm, variation factors Ullrich, does the (El-Shershaby, A1 on genes class target depends of of HSFs range as A importance such class the protection transport of proteotoxic confirm and diversification biosynthesis in background osmolyte proteins QK Liu as encoding in 2005, such genes pathways Schoffl, Salinas, & genes metabolical & Wunderlich different of (Busch, signaling Holuigue control, hundreds a Jimenez-Gomez, redox control forming (Olate, signaling, HSFs acclimation, pathways HSPs, cold A regulatory control Class pathogen to and 2018). reported cold (Qian recently integrates treatment were which responses peroxide hub pathogen and of changes regulator pH key (Bechtold heat, rape during oilseed age and Arabidopsis in tolerance 06 hn,L,Xn a,20) n a eue clmto aaiyt nxa(Banti anoxia to capacity acclimation reduced had and 2009), Gao, & Xing Li, Zhang, 2006, , sasrs-nue F hc otosrsosst etadsvrlohrsrse nmn plants many in stresses other several and heat to responses controls which TF, stress-induced a is tal. et 02.Oeepeso fHF2i Kmtn ol atal ecedevelopmental rescue partially could mutant QK in HSFA2 of Overexpression 2012). , 2 O 2 estvt u o h atadomtchprestvt,sgetn htti HSF this that suggesting hypersensitivity, osmotic and salt the not but sensitivity hsfa2 uatwshprestv oha n xdtv tess(Nishizawa stresses oxidative and heat to hypersensitive was mutant tal. et tal. et 01.Rdcdatvto fms etidcdHFgenes HSF heat-induced most of activation Reduced 2011). , 98.Oeepeso fHF2i rbdpi could Arabidopsis in HSFA2 of Overexpression 1998). , tal. et 3 03.HF1 ol iiihoiaiedam- oxidative diminish could HSFA1A 2013). , tal. et 04.HF1fcosadNR,a NPR1, and factors HSFA1 2014). , tal. et tal. et tal. et 06.Fnto fHSFA2 of Function 2006). , athsfa2 07.HSFA2-controlled 2007). , 05 Nishizawa 2015, , rbdpi mutant Arabidopsis tal. et tal. et tal. et 2017). , 2002). , 2010, , tal. et tal. et et Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. ruh n attlrneo rbdpi yicesn B estvt n peuaigmn B or ABA many and upregulating 2015), and McIntyre, 2020). sensitivity & Liu, ABA Drenth & increasing (Xue, Li by genes Zhao, Arabidopsis chaperon (Bi, of and genes tolerance HSP stress-induced of salt expression and enhanced drought through genes ABA-responsive wheat many of activated (Huang genes and Akhter, ABA-induced hypersensitivity Son, and ABA Jeong, Woo, increased Kim, HSFA6A Hwang, (Hwang 2014). 2016, Jinn, Bahk, & & Yang Choi Niu, to (Huang, hub plants a Arabidopsis salt-induced overexpressing as transcriptional and through functions homeostasis 5). factors drought HSFA4 cellular (Figure The that regulate genes and suggest, defense-related stimuli and data biotic protective These and of abiotic 2020). control of Zhuo, signals & & ROS Han Martinoia transmit Wu, An, Qiao, accumulating Liu, Choi, metal Zhang, Lee, the the from of Lee, SaHSFA4C hypersensitivity while Hwang, Cd 2016), (Shim, Huang, & expression Yang Zhuang, from (MT) 2017). PvHSFA4A Wang, metallothionein & 2009). enhancing Li Lee, Xue, by Liu, and rice (Lang, seeds to oligosaccharides osmoprotectant Arabidopsis type transgenic cadmium-induced raffinose Chen, The of of & tolerance biosynthesis Chen for desiccation Jiang, sible Song, increased Gao, of rape Zhao, sensitivity of oilseed Zhang, rescued tolerance (Li, class from CAT) salt BnHSFA4A A4 APX, Enhanced (SOS1, (SOD, 2007). transporters conditions. 2018). enzymes ion Nover, stress scavenger key various & ROS of and to Scharf activation HKT2) with responses Chan, correlated regulate (Baniwal, plants also chrysanthemum death overexpressing can cell CmHSFA4A plants and other signalling from ROS HSFs in action (Andrasi HSFA4A promoters their tuning to metabolic binding and directly scavenging by Perez-Salamo WRKY30 ROS or in ZAT12 al. enzymes as et such proteins, TFs defense-related defense-related HSPs, or response encoding genes hand (Farago other of heat the upregulation On to 2014). not Szabados, but & Lumbr- Rigo, Koncz Vilela, (Andrasi, Zsigmond, Bogre, Szabados, Medzihradszky, Rigo, 1) the & Darula, Papdi, (Figure Perez-Salamo, Domoki, Cseplo 2018, Horvath, damage combination Dasari, Nagy, Szabados, oxidative to & eras, Ayaydin, and Andrasi and Valkai, Baba, accumulation Arabidopsis anoxia Sass, Pettko-Szandtner, ROS Farago, in paraquat, Klement, 2019, reducing overexpression by Papdi, by HSFA4A generated Perez-Salamo, stresses stress Zsigmond, heat S1). oxidative (Table and cadmium, salt species salt, plant of to several tolerance in enhanced stresses oxidative and He metals proline Li, compromised Cao, to Zhong, (Wu due pathway Zhao, HSFA4 maybe Wang, catabolic plants Liang, the overexpressing Wu, LiHSFA3 of 2013, of activation sensitivity Li, by salt & accumulation Increased Xu Wang, 2018). Zhang, Yi, (Li, & ( hypersensitivity salt lily to or Koskull-Doring, lead von tomato Lim, & Hwang, from Vierling HSFA3 (Chen, Englich, connected of Ganguli, AP2- is Kiehlmann, the signaling by Larkindale, controlled osmotic Yoshida Schramm, is and 2008, 2010, HSFA3 heat of Lim, induction that & (Zhu, heat showing Lee levels and TFs, Kim, ROS Drought enhance DREB2C reducing 2020). also or and Lv, DREB2A contents could & type OsHSFA3 polyamine Zhou and Tang, rice ABA Zhou, of Gao, increasing Zhang, expression by Kidokoro, Arabidopsis Ectopic Mizoi, in Qin, tolerance 2008). drought Maruyama, Yamaguchi-Shinozaki, Todaka, Hin- & Sakuma, (Prandl, Yoshida, thermotolerance Shinozaki enhanced 1998, Fujita, and Schoffl, genes & chaperon Eggers-Schumacher and derhofer, HSP many induced HSFA3 Arabidopsis the HSFA3 2016). Baurle, & Altmann Brzezinka, hsfa4a 09 altv,Rzsy in,Segin,Oie,Cuu hle,Shac ite,2005, Mittler, & Schlauch Shulaev, Coutu, Oliver, Shengqiang, Liang, Rizhsky, Davletova, 2019, , tal. et tp atr oto epne ovrostpso tesssc sdscain ihlgt at heavy salt, light, high desiccation, as such stresses of types various to responses control factors -type euae aa hrooeac n ruh n aiiyrsoss(al 1.Oeepeso of Overexpression S1). (Table responses salinity and drought and thermotolerance basal regulates rbdpi uatwsfudt ehprestv oslnt,cdimadoiaiestress, oxidative and cadmium salinity, to hypersensitive be to found was mutant Arabidopsis tal. et tal. et 04.HF6 induced HSFA6B 2014). , 04.HF5wsfudt erpesro SAAfrightroioesadfine and heterooligomers forming HSFA4A of repressor be to found was HSFA5 2014). , 08.HF3fo te lnsmyhv ieetfntos osiuieexpression Constitutive functions. different have may plants other from HSFA3 2008). , athsfa4a TaHSFA4A tal. et athsfa4c aplmvaginatum Paspalum ed yidcn h aatnlsnhs genes synthase galactinol the inducing by seeds SAAadHSFA6B and HSFA6A iimlongiflorum Lilium tal. et 08 Perez-Salamo 2018, , uatb nuigRSsaegn nye Ce,Y,L,Wn,Lu, Wang, Li, Yu, (Chen, enzymes ROS-scavenging inducing by mutant and OsHSFA4A 06.Aogohrseis aSAFipoe thermotolerance improved TaHSFA6F species, other Among 2016). , DREB2A HSFA7B ybnigt t rmtradurgltdasto stress of set a upregulated and promoter its to binding by rmwetadrc ofre dtlrnet es and yeast to tolerance Cd conferred rice and wheat from nacdC oeac nyat(hn hn a,Liu, Tan, Chen, (Chen, yeast in tolerance Cd enhanced ofre hrooeac oAaiosspat,but plants, Arabidopsis to thermotolerance conferred ) 4 tal. et tal. et atr nacdsl n ruh oeac in tolerance drought and salt enhanced factors ol rnatvt aytre ybnigto binding by target many transactivate could 04.Srs oeac a soitdby associated was tolerance Stress 2014). , 2018). , eu alfredii Sedum GolS1 and GolS2 complemented respon- , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. lnsdfn hmevsaantptoei irbsb ohsiae muesse odtc the detect to system immune sophisticated a by microbes pathogenic to against targeted themselves 2014). be defend Honys, to pests & Plants and Solcova shown Gibalova, pathogens was (Renak, to AtREN1 biosynthesis responses with rRNA Defense similarity in temperatures. 2.2 high implicated high has is Arabidopsis to it that in hypersensitive suggesting AtREN1 is nucleolus, 2015). which Chen, germination & The pollen Shen development. pollen to Zhao, regulates Wang, important but HSFA5 Lang, is Hou, module PeHSF-PeWRKY1 Sa, of that Zhao, character suggesting extremophile tolerance, the salt determine improved of tobacco elements in HSE PeWRKY1 the to binds (Ma, HSF was genes Other resistance induced pathogen and stress 2013). tolerance of stress S., osmotic number (Peng thermotolerance, reduced a basal upregulating while tobacco, by tolerance overexpressing enhanced stress-induced Arabidopsis of to B-type, in lead The chickpea drought of and 2015). CarHSFB2 stress-induced temperatures the high of expression to Ectopic the 2017). to Schleiff, attributed Bharti, & were Koskull-Doring, functions Von activator and (Bharti, Repressor Kay, stress stress-induced 2004). & during Nover, Pruneda-Paz genes & Treuter housekeeping Chow, Tintschl-Korbitzer, of (Kolmos, Kumar, expression stress maintain ( heat to tomato and contribute wild salt HSFB2B Response from flowering. moderate Pseudo HSFB1 late of during and 2014). promoter rhythm elongation the hypocotyl Bagman, circadian to to binds the Iwase, leading which transcription regulates functions signals to Rymen, its stress HSFB2B repressing stimuli abiotic Arabidopsis Shibata, (PRR7), with 7 The upstream (Ikeuchi, clock Regulator circadian is 2018). connecting responses of HSFA2 Sugimoto, promoter integrator wound & with an HSFA2 Brady as together of to Ahnert, HSFB1 Takahashi, control Favero, bind Arabidopsis Coleman, transcriptional to Watt, 2011). the shown Ohme-Takagi, was in & HSFB1 implicated Mitsuda (Ikeda, HSFs. targets A downstream desiccation class for important of is action and embryogenesis late of HSFB oxida- regulator from key 2012). seeds. apparatus a Jordano, drying is photosynthetic & of HSFA9 Diaz-Espejo the tolerance that Lindahl, protect suggest Tejedor-Cano, and results Personat, plants These Prieto-Dapena, heat tobacco dehydration, (Almoguera, to transgenic Mer- damage tolerance tive of Personat, confer could Almoguera, stresses HaHSFA9 genes of oxidative Prieto-Dapena, overexpression target Constitutive and 2020, CRY and 2017). Jordano, Jordano, PHY & & of Ruiz chan transcription Carranco, promote Prieto-Dapena, and signaling (Personat (Almoguera, light genes Tejedor-Cano blue HSP-coding 2006, enhance of photomorphogenesis, Jordano, tolerance expression & dehydration the Almoguera improved Together enhancing and Castano, by 2009). longevity seeds Jordano, seed tobacco enhanced & HaHSFA9 in Carranco of Espinosa, overexpression HaHSFA4a, Diaz-Martin, with Transcrip- Prieto-Dapena, (Almoguera, 2010). 2008, dehydration Jordano, Jordano, to & & Tejedor- Ohme-Takagi Almoguera of Hiratsu, Personat, Castano, tion Carranco, 2002, Prieto-Dapena, Almoguera, 2014, Jordano, and Prieto-Dapena, Jordano, & seeds Tejedor-Cano, & Carranco of Koskull-Doring, Almoguera ageing Prieto-Dapena, von and Prieto-Dapena, dehydration & Diaz-Martin, Cano, temperatures, Baumlein Rojas, high Vierling, to (Almoguera, (Kotak, tolerance embryos regulates maturation sunflower seed in HaHSFA9 during of 2007). protection Expression cellular to desiccation. tributing seed and embryogenesis abi3 late controls 2019). Wang, which & ROS TF Liu reduced Zhang, homeostasis, Wang, ion (Zang, HSFA9 stabilized potential through osmotic tolerance of salt adjustment enhanced and to accumulation leading motifs promoter E-box uat hl B3oeepeso nacdHF9epeso,wihcnrl eea S ee con- genes HSP several controls which expression, HSFA9 enhanced overexpression ABI3 while mutant, HaHSFA9 tp atr r osdrda rncitoa ersoso ekatvtr hc ouaethe modulate which activators weak or repressors transcriptional as considered are factors -type sase pcfi Fi eea lns rbdpi SA srgltdb B3 h edspecific seed the ABI3, by regulated is HSFA9 Arabidopsis plants. several in TF specific seed a is ee nldstesalt-induced the includes genes SlHSFB1 a nue yHDE2 nA2tp F hc otosgn xrsini response in expression gene controls which TFs AP2-type an HaDREB2, by induced was ftmt,wihwssont nac tesrcvr Rt,Mrs ulk Scharf Bublak, Mirus, (Roth, recovery stress enhance to shown was which tomato, of PeWRKY1 yoescnperuvianum Lycopersicon VpHsf1 .euphratica P. rmcieewl rpvn nacdaqie thermotolerance acquired enhanced grapevine wild chinese from rmtr n rmtsissl nuto.Oeepeso of Overexpression induction. salt its promotes and promoter, PeHSF atren1 5 tal. et uathsanra al olnmtrto,reduced maturation, pollen early abnormal has mutant rmtedsr olr( poplar desert the from Se,Yo u,Cag ag ig in Zhang, Qian, Ding, Wang, Chang, Sun, Yao, (Shen, 00.HHF9wsas hw opromote to shown also was HaHSFA9 2010). , ol c sc-ciao fcasAHF and HSFs A class of co-activator as act could ) HSFA9 tal. et ouu euphratica) Populus 04 Prieto-Dapena, 2014, , scmrmsdi the in compromised is which , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. eunl eerse SA.TeHF2RF ouerdcdtsRAboeei hc nacdthe enhanced sub- which which biogenesis (BRM), tasiRNA BRAHMA RELATIVE reduced factor demethylase module histone remodeling HSFA2-REF6 the chromatin The medi- induced the HSFA2. HSFA2 factors and derepressed temperatures HSFA1 (REF6) sequently high 6 In 2018). FLOWERING van EARLY Rhodes, 2017). Baurle, Ravarani, OF genes, Wigge, & Buning, target & Brestovitsky, Charng from Jaeger Charoensawan, nucleosomes (Cortijo, Liu, Noort, H2A.Z activation Hung, of large-scale removal their Lin, by transcriptional triggering necessary rearrangement, Lamke, fast chromatin was the (Liu, temperature-dependent facilitated which genes ate which methylation memory, target K9ac stress of of (Lamke and activation regulation exposures tri-methylation epigenetic micro HSFA2-dependent HS and involves 2014). the Baurle, (Charng regulation, repeated di- ence & ago epigenetic after Asso- H3K4 Lamke Brzezinka, decade on hyper-induction promote (Stief, known. relies a for to promoters well memory than target found not of stress more was methylation is such described histone HSFA2 regulation that of been modulation revealed, epigenetic has and data in memory RNAs recent role stress More their with responses, 2007). HSFs stress several in of and essential ciation HR are for HSFs essential While regulation is in These epigenetic pathogens which in various 2018). signals involved to Yu, HSFs ROS resistance & 2.3 enhancing regulate Zhou and can Shi, coordinating HSFs Xia, B by resistance. Yin, class mainly disease Cao, and species, Xu, A In plant (Zhou, class HR. different HR several to that via leading Nematode demonstrate, genes defense HSFA-type results 2019). H basal of Chen, apoplastic activation and are for & and accumulation essential HSFs Geng burts be HSP90 Xia, various ROS to Li, to found that Hou, was leads SlHSFA1a suggesting Hong, roots tomato Yao, mealybugs, (Yu, tomato plant resistant and this of disease, in infection 2018). resistance streak Shi, disease & brown in He cassava implicated Chang, blight, Liu, (Wei, bacterial MeHSF3 disease of cassava A targets in Pathogen-Related direct class resulted and be to MeHSFA9B (EDS1) to MeHSF3 MeHSFA6A, 1 belongs turned of which Susceptibility cassava (PR4), silencing Disease of 4 Enhanced VIGS-mediated gene MeHSF3 of expression The infection. reduced Xanthomonas and 2016). hypersensitivity to Minami, responds & which elicitor-induced Nishizawa HSFs Shibuya, the Toki, with Hagio, Together Fujisawa, 2002). OsHSF23 Yamada, & The Ashikari OsHSF23 also. 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DREB2C Lim, downregulate & to with identified Hong interacting recently was by (VOZ1) expression 1 HSFA3 ONE-ZINC-FINGER HSFA3 PLANT (Chen VASCULAR induce stress factor Schramm could heat domain 2006, during DREB2C Yamaguchi-Shinozaki, expression & and of Shinozaki DREB2A level Osakabe, high for 2011). needed Yamaguchi-Shinozaki, AP2-type Osakabe, were Todaka, & and considerably the Seki, Shinozaki Kim, as or Maruyama, Schoffl, such Nakashima, abolished Mizoi, Sakuma, TFs, Kidokoro, was of Nakajima, other genes Ohama, levels of (Yoshida, HSF mutant transcript activation activate heat-induced while Heat-induced and mutant, most released QK 3). of are the activation HSFA1s in that stress, reduced (Scharf showing heat HSFs proteins data, Upon heat-induced the transcript and the chaperons. genes including expressing HSP70/90 genes, constitutively with by downstream encoded complexes are inactive factors HSFA1 form upregulated. the also were while genes treatments, stress HSFB2A any HSFB1, S4). eFP by HSFA7B, S3, to induced (Table According weakly ecotype treatments Col-0 only stresses. more In were biotic or genes and one to abiotic HSF different responsive Arabidopsis were to 21 remaining response the considerable in of revealed genes 7 databases HSF Browser, Genevestigator individual or the Browser of S2). eFP (Table variation Arabidopsis climates from warm extracted in data Transcript Comparing collected conditions ecotypes environmental ecotype. in extreme laboratory higher to standard were Responses the genes 3.2 HSFA1 Col-0, of including levels climates transcript continental Col-0, mild to from plants in low hsfa7a 04 Swindell 2014, , ol nac h xrsinof expression the enhance could ee.Wiethe While genes. while , tal. et uat,sgetn htteeHF euaectslcpoenrsos Ln si u u& Wu Lu, Tsai, (Lin, response protein cytosolic regulate HSFs these that suggesting mutants, HSFA7A 07.Tasrp eeso hs ee eeifro nthe in inferior were genes these of levels Transcript 2007). , etstress heat HSFA7A SA,HF4,HF6,HSFC1 HSFA6B, HSFA4A, HSFA2, tal. et and rcf2-1 a oneuae.Wt h xeto of exception the With downregulated. was 07.Sm fteeHF eehgl nue ysl nthe in salt by induced highly were HSFs these of Some 2007). , ol activate could and nue h xrsino rbdpi S genes: HSF Arabidopsis 7 of expression the induced HSFB4 and HSFB2B nac019 eesprse Fgr ,TbeS) rncitpoln aacon- data profiling Transcript S3). 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All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. S ee hog D1 A4adS-eedn inl.Ifcino Seoyeby ecotype WS of Infection & signals. 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S3), 2002, Fernie, Kobayashi, & (table Fukushima, Saito minutes Tohge, Matsui, within Kusano, Niida, Matsuda, 1997, Kawashima, Khurana, Goto, & of (Bharti sets expression similar stimulated of ozone and Induction the UV-B 2007). 3). , (Figure S3). heat table heat: by by 3, repressed induced but were which light genes genes: excess HSF by same photosynthesis HSFB2A induced the disrupted was HSFB1, induced indicating HSFA7B, light genes, 3). HSFA7A, Strong HSF (Figure HSFA3, changes of 3). HSFA2, transcript number (Figure the a response of of stress all expression generates not the but enhanced some darkness reversed conditions Sustained hypoxic from Recovery conditions. different induce (Nagaraju to sorghum shown as were such stresses Such plants and salinity. signals other Xue ABA-dependent and by in drought controlled during genes are TFs HSFs HSF Arabidopsis ABF-type of and subset DREB a of that MYB, promoter suggest the results to These bind 2016). could HSFA6B activate hand, could other TF the ABF2 the while tant, S rncitpolsi rbdpi.Fs nuto of induction Fast Arabidopsis. of in upregulation profiles transcript HSF Marriott, 2013). Temanni, Toneva, stresses Hille, & Minkov, Other Fernie Sujeeth, Schippers, Tohge, (Zhang Mueller-Roeber, Obata, respectively Antonio, Benina, (Gechev, Thomas-Oates, stress, survival Bergstrom, drought for (Fragkostefanakis and important responses salt are they stress by induced of regulator be plant resurrection key can as genes defined HSF HSFA2 twelve and al. stresses et many (Guo stresses by both induced by upregulated were were genes three (Chauhan while respectively, drought and cold by 04,i ha n ie(Shim rice and wheat in 2014), , sng1-1 tal. et 05.Gnm-ietasrp nlsso hscnt(arpacra . eeldta i and six that revealed L.) curcas (Jatropha nut physic of analysis transcript Genome-wide 2015). , SA,HF4,HF8 HSFB1 HSFA8, HSFA4A, HSFA2, , SBA HSFB2B HSFB2A, 04,o iei hc sSA epne togyt et sSA a tmltdmostly stimulated was OsHSFA3 heat, to strongly responded OsHSFA2 which in rice or 2014), , uat(iue3.SG otostepeypoaodptwyadteU--eedn gene UV-B-dependent the and pathway phenylpropanoid the controls SNG1 3). 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All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. ihrdcdH reduced with & of Mullineaux Expression Panchuk, (Volkov, 2007). antioxidants by blocked be (Perez-Salamo could stress 2006). heat Expression Schoffl, during 2015). genes Blumwald, (APX) & num- oxidase Mittler a H 2017, induce 4). 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(He transcript out not NMD and is MsHSF1, genome function of by their mechanism isoforms exceptions identified splicing five few been with specific have species, variants a plant splice many that HSF 2018). suggesting other Cho, numerous temperatures, & Although high Yoo alternative (Kim, in mediate conditions transcripts (Liu to such HSP site reported at and splice exist and mRNAs identified HSF 5’ degrades was of intron (STA1) which splicing cryptic 1 mechanism a STABILIZED a through protein, NMD, U5-snRNP-interacting formed by was eliminated HSFA2-III (Sugio is of codon which termination HSF form, premature Arabidopsis HSFA2-II with most inactive Schleiff for the Scharf, (Liu indicated erating Bovy, Zou, poplar is Bublak, Xie, transcript 2012), Gebhardt, (He, single Kosuge, and database alfalfa Roth, & for TAIR 2009), transcripts Jimenez-Gomez, Iida the two Maule, Mesihovic, (Amano, In genes, Arabidopsis & (Hu, species Aparicio in 2020). tomato pondweed Fragkostefanakis, Dreos, reported in & aquatic Sugio, been and in 2013, also 2019a) 2007), Qi, have , lev- Yu, & transcript Wang (NMD) & of Du, Zhu decay modulation Liu, Wang, nonsense-mediated regulation, Liu, and HSF Sun, splicing in (Liu, mechanism alternative dominant by the els is control transcriptional HSFs While plant of (Driedonks regulation genes transcriptional stress Post target 3.5 other downregulate and ROS HSFs 2015). attenuate of can targets signals antioxidants direct ROS be dependent can While CAT or 2012). 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Aro, as & such genes Kangasjarvi Lohrig, scavenging Suorsa, (Jung Pesaresi, ROS Gollan, (Ihnatowicz, HSFs, Tikkanen, stress-induced nuclei activate 2008, and can is chloroplast Leister, between and pool & signaling senescence phos- Muller plastoquinon retrograde homeostasis, Wolters, kinase of redox of STN7 overreduction regulates component The prevent proteins, principal 2004). antenna a to Leister, harvesting and & light flow Salamini photosynthetic Jahns, electron phorylates Schneider, ROS sustain Richly, Varotto, these to Pesaresi, between important (Ihnatowicz, had crosstalk is genes a the HSF protein ROS-induced is in (PSAD1) there 2007). levels Apel, peroxide, transcript & hydrogen Murgia reduced and Warzych, Pers-Kamczyc, oxygen Stachowiak, singlet (Laloi, and pathways superoxide, Hideg, and chain 2010) by Wagner, transport Padmasree, induced Danon, electron & are Raghavendra mitochondrial Kim, Yearla, the Abhaypratap, Laloi, blocks (Dinakar, A superoxide Ochsenbein, upregulated Antimycin toxic Przybyla, highly 2003). the Camp, Apel, generate den & Nater (op Feussner, 4) Gobel, (Figure damage oxidative HSFB2B 06.RSsgaigtruh Sshsteeoeafebc euaoymcaimb hc HSF- which by mechanism regulatory feedback a therefore has HSFs throught signaling ROS 2006). , HSFA4C tal. et eeidcdi the in induced were tal. et SA,HF4,HF8 SB,HSFB2A HSFB1, HSFA8, HSFA4A, HSFA3, 03 oao aaea eGr ePno 08 Perez-Salamo 2008, Pinto, De & Gara De Gadaleta, Locato, 2013, , wwaaiossog.Avrecniin a rmt lentv piigo SA gen- HSFA2 of splicing alternative promote can conditions Adverse (www.arabidopsis.org). 07.Tendl pcfi sS1mN a rdcdb rn-piigbetween trans-splicing by produced was mRNA MsHSF1 specific nodule The 2007). , tal. et 09) nwl oaoseisatraieslcn eutdi cuuainof accumulation in resulted splicing alternative species tomato wild In 2019a). , SAB HSFA1D HSFA1B, flu uat hrceie yehne ige xgnacmlto n photo- and accumulation oxygen singlet enhanced by characterized mutant, stn7-1 and tal. et psad1-1 and tal. et 09.I xrm ihtmeaue e pievariant splice new a temperatures high extreme In 2009). , HSFA3 tal. et 11 uat Fgr ) h htsse uui D-1 subunit I Photosystem The 4). 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Such provide and can complexes. location, interactions regulation intracellular protein-protein protein their regulate mapping or initiation, about Therefore transcription proteins for degradation. capacity of and their the number stability define networks, a and regulatory with HSF in interact HSFs the physically on can PROTEINS depending HSFs HSF activity, PLANT reduced OF or INTERACTIONS enhanced of 4. confer 2018). modifications Meng, translational can & post which frequent Ma modification. plants, Wang, are introduced in sumoylation Liu, enhanced, and Lv, proteins SlSIZ1 Wang, phosphorylation HSF of that (Zhang, of Overexpression revealed induction tolerance plants. data the heat heat-stressed promoting These reduced in tomato silencing accumulation was in ROS SlHSFA1 RNAi-mediated (SlSIZ1) with of of while ligase co-activation reduction sumoylation E3 synergistic and mediate SUMO genes and and SIZ1 HSP with was promoter The which interact target sunflower, 2017). to a of Jordano, shown HaHSFA9 of recently & specific activation Almoguera seed enhanced Prieto-Dapena, activate the Meiri, to (Carranco, on Schuster, HSFA4A to leading reported (Cohen-Peer, capability was K38, thermotolerance effect at its Different acquired sumoylated reduced of 2010). 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HSFA2 sumoylated SUMO1 plants that in and genes. demonstrated modification SUMO1 HSP reports translational with several post HSFs, important interact plant an could of be sumoylation can on conjugation Sun, available SUMO Chang, is Li, information HSFA1A (Liu, shock limited with responses heat Although HSFA1A stress interact mediate of heat could dephosphorylation promoted 2007). to PP7 PP7-mediated and Li, shown HSPs & system. was genes. target Zhou (Y2H) (PP7) of hybrid HSP two expression 7 of yeast enhanced phosphatase upregulation in subsequently CaM3 to protein calmodulin contribute binding the and calcium and Arabidopsis multiple The at in phosphorylated known. response be well can on it not (Andrasi that phosphorylated is suggesting kinases was targets, different kinase and by MAP MPK6 sites (Andrasi from different genes and residues target MPK4 acid of amino MPK3, transcription Koskull- with and Perez-Salamo von multimerisation interacted phosphorylated Lucks, 2019, intramolecular HSFA4A was Lecourieux, promoted and Kumar, which with (Evrard, 2013). Zhang, Ser309, interacted stress Zhang, Hirt, HSFA2 heat Su, Arabidopsis & biotic during 2012, The Mundy, various transfer Doring nuclear & to 2017). Petersen 2002). facilitating Zhang, responses Roux, MPK6, Roitsch, & accelerate by Rasmussen, & Xu 2018, and Lukowitz, Ehness kinase, Hirt, signaling, Liu, MAP & Proels, ROS Sun, (Bigeard calcium-dependent Hofmann, mediate stresses a Vashista, MPK6 abiotic of Sinha, in and and substrate 2008). HSFs (Link, MPK4 a Zhou, plant responses MPK3, is the & stress Arabidopsis tomato of Sun some heat in Liu, HSFA3 phosphorylate regulates Han, can heat-induced which Li, which The & Gao, transduction (Liu, Koncz signal conditions. ROS thermotolerance transcription stress Schell, mediate enhanced basal Schoffl, (AtCBK3) kinases increased 3 MAP (Reindl, and kinase Several capacity genes protein HSP binding CaM-binding of the HSFA1A DNA by activation Arabidopsis reduced phosphorylation of while Phosphorylation kinase 1997), regulation. CDC2a Bako, such and cyclin-dependent in complex implicated more the is are phosphorylation by kinases through protein Sistonen, activities HSF of & of classes modulation several Johnson families, HSF Thiele, large have Denessiouk, plants reduce As Hietakangas, which 2017). (Anckar, Dinkova-Kostova, domain & activation regulatory Naidu the gene Stevenson Dayalan Morrice, on target 2006, Price, sumoylation Hellman, and Soncin, or Meinander, (Chu, 2001) binding Kallio, activity Sistonen, DNA Rantanen, reduce & Mikhailov, or Eriksson Hietakangas, rather phosphoryla- enhance Morimoto, a including MacKintosh, either Holmberg, in modifications can 1996, expressed translational which Calderwood, is post residues cells & by serine animal regulated multiple in is at HSF1 of activity tion activation 2004). its the Sakurai, and for & manner (Hashikawa prerequisite constitutive HSEs was atypical HSF yeast with of genes Phosphorylation target sumoylation. or acetylation phorylation, tal. et 04.Ms pcrmtyaayi fHF4 ol dniyphosphorylated identify could HSFA4A of analysis spectrometry Mass 2014). , tal. et 09.Rl fpoendpopoyaini euaino HSFs of regulation in dephosphorylation protein of Role 2019). , 12 hsfa2 uatwt eue xrsinof expression reduced with mutant tal. et , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. eeoesbtencasAadBHF a hrfr efre npat ersnigrfie transcriptional refined representing plants in of formed combinations genes. be Multiple therefore target Singh, can 2012). of Lavania, HSFs Grover, control B & Enoki, and Mishra (Mittal, A with Agarwal, class Lavania, confirmed OsHSFA7 between Mittal, be heteromers Singh, OsHSFB4b, could 2011, with Grover, OsHSF26 & OsHSFA2a and Sakurai OsHSFB4c of with to heteromerisation OsHSFB4b lead and OsHSFB4b, CBP/HAC1 OsHSFB4b induction of gene OsHSFA9, for Suppression essential OsHSFA2c, is activation. (CBP) complex ternary protein synergistic such binding that (Bharti to suggesting CREB leading activation, plant transcriptional promoters the of ( abolishment target with tomato complex on wild ternary HAC1 enhanceosome-like of ortholog formation an LpHSFB1 formed trimer LpHSFA1 genes. LpHSFB1 factor heteromeric target and A of or class expression with homo interact the containscould for tune (Li OD sufficient fine HSFs conserved can and B The class factors required and HSFB2B. are A and class which HSFB1 between repeats Arabidopsis (Tejedor- heptad for HSFs hydrophobic reported these 2014). were Jordano, between interactions & synergism Homomeric Espinosa transcriptional system Almoguera, Y2H Prieto-Dapena, promoting in Personat, HSFA4A detected Carranco, of Cano, be was retention not HSFs type could nuclear HSFA5 A caused and class HSFA1A other between with interaction activation (Baniwal and Gene HSFA5 signaling. by can ROS HSFA4A affected in (Baniwal not While activation module HSFA4/A5 gene HSFA5. the such of of represses OD HSFA5 conservation through with interaction tomato al. genes, and bonds stress-responsive et Arabidopsis of Cys-Cys in number (Andrasi formed promoting a interactions induce be and intramolecular can residues HSFA5 such by and Cys enhanced influenced HSFA4 conserved was their MPK6 HSFA4A of to or Arabidopsis oxidation due MPK4 of by sensors Trimerisation stabilised peroxide was 2006). (Perez-Salamo as which Mittler, activation function the status & facilitating to environment, redox (Miller oxidative factors to genes response these ROS-induced in heterotrimers enable of or suggested homo active complexes was form to superactivator HSFs capacity of hexameric of transport in sensitivity nuclear resulting 2009). ROS for Scharf, homomers essential & to Nover was preferred regulatory Bublak, HSFA2 was (Scharf Baniwal, overlapping and (Chan-Schaminet, activation oligomers 2010a). in HSFA1 transcriptional hetero Schoffl, of function HSFA2 synergistic & interaction they and to Berendzen tomato, that leads (Li, In which suggesting genes HSFA2, HSFA3, target 2017). with certain (Li, also of HSFA2pathway late induction interacts the HSFA2 synergistic with HSFA1B showing Arabidopsis HSFA1A, early reported between also interactions Schoffl, was Heteromeric heterotrimers & way. target and Berendzen redundant Oecking, homo in Weckermann, the form Doll, response in other, (Li, each genes HSEs Yoshida target with common interact 2010b, to activate HSFA1D and binding and combinations HSFA1B all HSFA1A, for in species. required plant is 5 in which Scharf heterotrimers, 1999, and Nelson, homo (Lee form promoters factors shock HSFs. plant Heat of interactions heteromeric cases and 34 techniques. Homomeric in more method, 4.1 one or with two and detected by Thellungiella were confirmed coupled interactions 8 lily, were methods protein-protein as interactions in most capture soybean, such affinity While of sunflower, methods, or spectrometry. several pull-down interactions rice, mass with complementation, with 35 maize, fluorescence demonstrated bimolecular were and tomato, hybrids, interactions two HSFs, wild Protein-protein yeast Arabidopsis were and 1). (Table interactions of cultivated species protein-protein interactions as Eriobotrya 221 186 such Altogether including species databases. proteins other public HSF and with papers identified scientific from data imental 07.Tefc htsmlratvto/ersineiti neae ln pce ugssfunctional suggests species plant unrelated in exist activation/repression similar that fact The 2007). , tal. et tal. et 04.Oioeiaino Sswsosre nrc lo hr ooe omto of formation homomer where also, rice in observed was HSFs of Oligomerisation 2004). , tal. et tal. et tal. et 07.I uflwrHHF4 ol neatwt h edseicHHF9which HaHSFA9 specific seed the with interact could HaHSFA4A sunflower In 2007). , tal. et 95 ilr&Mtlr 06 eeadr,Rbnti,Si,Lu emr ig& King Wemmer, Liu, Shin, Rabenstein, Peteranderl, 2006, Mittler, & Miller 1995, , 04.Popoyaino h osre e39rsdeb A iae MPK3, kinases MAP by residue Ser309 conserved the of Phosphorylation 2014). , 01.Mlil neatoscnr htcasAHF euaeeryha shock heat early regulate HSFs A class that confirm interactions Multiple 2011). , 02.1 ooei n 6htrmrcHFHFitrcin eedetected were interactions HSF-HSF heteromeric 26 and homomeric 10 2012). , tal. et nvivo in 00,Peteranderl 2010b, , n ucinda ociao ftre ee.LpHSFA1 genes. target of coactivator as functioned and 13 tal. et tal. et 99.Itrcino ls n B and A class of Interaction 1999). , tal. et 98.Fraino HSFA1 of Formation 1998). , 09.Htroioesof Heterooligomers 2019). , yoescnperuvianum Lycopersicon ) Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. ciiyo tblt Si,Jz ar,Zag&Shcta,21) eetyitrcino Arabidopsis of interaction Recently 2011). were HSFA8 Schachtman, and & HSFA1A HSF Zhang influence may Basra, proteins. 14-3-3s cytoplasmic Jez, that suggesting other (Shin, proteins, with stability binding interact 14-3-3 or the to activity among reported analysis MS/MS were by HSFs identified corepressor as the function of and with Tripp, HSFs Some together various (Port, that with HSFA2 suggest complexes suppressed chaperon results control. form which These transcription can tomato, in HSPs 2004). of small Scharf, HSP90, cytosol & and the Bublak HSP70 in Winkelhaus, HSFA2 Heerklotz, Weber, with demonstrated Zielinski, aggregates was large transcript 2011). factors Scharf, formed OsHsfA2c regulated & and (Singh CII HSP90 Schleiff tests OsHsfB4b BiFC Bublak, with HSP90. and (Hahn, (ClpB-cyt) Y1H by HSFB1 HSP100 in enhanced of cytoplasmic was degradation rice HSFA1, of proteasomal HSFB1 of Interaction promoted of activities and the binding HSFA2 repressed for of DNA needed and levels while isomerase, with cis/trans interacted HSFB1, prolyl HSP70 and peptidyl Tomato HSFA2 a thermotolerance. is of ROF1 prolongation 2009). the Breiman, & (Meiri domain. complex phosphorylation (TDR) ROF1-HSP90-1 HSF repression by promoted temperature-dependent was the A which (Higashi temperatures at (Ohama Class high place responses in several took dissociated shock with complex with and HSFA1D/HSP70 heat HSP90 The interact activation or blocking could the HSP70 salsuginea, HSP90-3 with by Thellungiella interaction and mediated of HSP90-1 was ThHSFA1D al. HSFA1A 2002). and et Arabidopsis Schoffl, Arabidopsis plant with & of several HSP70 in (Kim HSFs HSFs of domain various and binding Interaction chaperons with DNA HSP90 1). or Sistonen, complex HSP70 (Table & between in Morimoto identified species (Akerfelt, cytoplasm were trimers interactions active the Scharf 23 is form 2018, complex in 1998). to Thiele, the nuclei & conditions retained the Burchfiel stress into Gomez-Pastor, are In transported 2010, activation. HSFs are transcriptional HSFs inactive from and HSFs dissociated model, prevents which cycle identi- HSP90, were and shock activation HSP70 proteins heat the HSBP A two to class maize several According 2006). In with Scanlon, interaction 2010). & non-redundant thermotolerance Nover Jinn, Rogowsky, showed acquired and & (Fu, which HSFA1B promoted HSBP Lai factors HSFA1A, HSBP2) but two (Hsu, and with development or response (HSBP1/EMP2 HSBP seed one shock fied Arabidopsis for have heat essential Plants the and and was between binding binding 1998). and DNA Interactions promoter Morimoto, reduces target & genomes. HSF1, regu- reduced Kramer trimeric their HSFA2 negative Fox, dissociates in are Chen, HSBP1 which genes (Satyal, cells proteins, coding genes animal nuclear target In small of conserved response. activation shock are heat (HSBP) of proteins lators (Czarnecka-Verner binding domain model factor prevents repressor however shock Drosophila the (GmHSFB1) Heat the via soybean from to TFIIB HSF with analogy B during interaction Class In reported via A was transcription activation. 1998). TBP2) transcription promote Schoffl, (TBP1, therefore & 2 and (Reindl and 1 promoter HSFA1A protein HSE-containing (Jedlicka Arabidopsis binding of a TATA Interaction complex of factors SWI/SNF TFs. the recognition transcription of with SWI/SNF binding general associated the DNA be the can modulate of HSFs with and the components Hong, structure of Birnbaum, several chromatin some Steiner, that regulate with Wu, suggest which Kim, Results interact Han, 2013). can (Efroni, Wagner, SWI3C HSFA6A & and Eshed and SWI3B 1). HSFA3 BRM, (Table complex, that remodeling protein revealed chromatin domain Homeobox, screen ZnF, bZIP include or Y2H and bHLH A families, ERF, protein Zhang, MADS-box, different Huang, other NAC, to by O’Malley, belong VP1-B3, Feeney, validated TCP, TFs Goubil, been HSF-interacting MYB, detected not Castanon, 2017). however have the Ecker, Bartlett, were and of & Nery, Arabidopsis interactions category Galli MacWilliams, of such common Garza, mapping most Most (Trigg, interactome The CrY2H-seq methods interactions). proteins. throughput (69 high other TFs a of in range other a were with proteins interact HSF-interacting can HSFs trimerisation, Besides proteins other with Interactions 4.2 03 Yoshida 2013, , tal. et tal. et 97,patHF1cnfcltt idn fTP oTT o hog ietinteraction direct through box TATA to TBPs of binding facilitate can HSFA1 plant 1997), , 06.HF2cuditrc ihHSP90-1 with interact could HSFA2 2016). , tal. et tal. et 01.Tasciaincpct fAaiossHF1 a eue by reduced was HSFA1D Arabidopsis of capacity Transactivation 2011). , 02.SvrlsalHP a loitrc ihpatHF.HSP17.4- HSFs. plant with interact also can HSPs small Several 2012). , 14 tal. et 02 o,Go utoce mt Voellmy, & Smith Guettouche, Guo, Zou, 2012, , nvivo in uprigncertasoto the of transport nuclear supporting , tal. et 2004). , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. ru rti HG)fml,rqie o N earadrgltsrsosst eooi agents is genotoxic protein to TIM-barrel The responses regulates 2018). Amiard, and & repair Butter DNA Mobility White, High Benyahya, for the a Ines, required to is Da belongs family, (REC1) Rymarenko, HON4 Sinkler, (HMGA) 1 protein (Charbonnel, Stavoe, coverage histone A Ruckle, linker chloroplast Protein Stefano, The Reduced (Larkin, Group 2016). nuclear development Osteryoung, The GTPase chloroplast & mitochondrial Malmstrom & regulates N-terminal 2006). Kawamura Brandizzi, with which Kondo, Innes, an signaling protein Furuto, proteins. & SA Tominaga, with binding Frye regulatory (Minami, linking mRNA dynamins Ade, tolerance diseases these Tang, freezing to fungal of in 2015, to related implicated Uemura, functions resistance was is control and apoptotic protein to PCD the shown functions, (EDR3) was and 3 EDR3 UPR gene, resistance domain. ER-based related disease between closely Enhanced link the is The that a BAG7 (Nishizawa-Yokoi (Li, intriguing, represent HSFA2 folding is 2010). may of protein of It responsible targets Dickman, control is stress-induced & through and 2017). the stresses TFs Britt biotic Dickman, with Kabbage, and interact & abiotic can Chen, Williams, Williams during it (Doukhanina, homeostasis where 2006, conditions cellular nuclei, Dickman, stress maintaining and for abiotic (ER) & reticulum in Reed endoplasmic or (UPR) between Godzik, attacks response shuttling pathogen protein Zalm, in unfolded der regulates (PCD) van mammalian and death the chaperons of cell and homolog HSP70 HSFA4A programmed plant with between a and is interacts formation (BAG7) which complex 7 athanogene regulator, facilitate BCL-2-associated apoptotic may The HSP90 HOP2, HOP2 and proteins. protein HSP90 HSP70 2018). (TPR) Fernandez-Calvino, or (Fernandez-Bautista, UV-B with Castellano, the HSP70 repeat stress interacts & and of genotoxic tetratricopeptide which Mock and heat component Toribio, The cytosol, thermotolerance by a Munoz, the acquired 2016). induced in is and Cao, implicated nuclei be PRP8 is & between can and factor shuttling Liu and both splicing co-chaperon Kong, splicing pre-mRNA as with Sun, functions in The Gu, heteromers involved Wang, and form cytosol. (snRNP), Lu, diverse can in ribonucleoprotein quite (Deng, HSFA4A were or nuclear proteins that nuclei small HSFA4A-interacting Piconese, Other in shows U5 (Fortunati, and HSFs. either results HSFA4A development other localized Our with to root be were not similar could and but is 2008). HSFA4C, which HSFA4C regulation and Migliaccio, HSF less-known HSFA5 hormonal interacting The & response, other repressor Ferrari samples. gravitropic the Tassone, as IP was in salt-treated acted HSFA4C implicated and HSFA5 was 2018). control which (Huang, both in Formation in irradiance demonstrated detected samples. high control already to which and sensitivity was proteins, salt-treated the HSFA4A in immunoprecipitated (Baniwal similar the and was HSFA4A among HSFA5 abundance of score its of highest and heterodimers samples the IP of had all HSFA5 Results in spectrometry. present 2. mass was with Table with experiments in identified (Co-IP) were summarised coimmunoprecipitation proteins are of immunoprecipitated series the out and (Andrasi HSFA4A carried YFP-tagged have kinases we MPK6 interactors, (Davletova HSFA4A and thermotolerance further not MPK4 but MPK3, responses, by stress oxidative and al. salt et controls are HSFA4A HSFs Arabidopsis class The function. B proteins silencing interacting As gene HSFA4A their of 2012). HSFs, enhance Identification Davies, various might 4.3 & recruit proteins DNA, to Guo TPL/TPR to shown Ashworth, with bind were interaction (Causier, directly corepressors repressors, HSFB2B TPL/TPR not transcriptional and does TFs. HSFB2A which other HSFB1, regulation, of including several function transcription repressing in identified enhance corepressors screen but as (WAMP) Y2H function protein proteins family (Li pathways. membrane (TPR) protein vesicle-associated regulatory calmodulin-binding a various a including protein-protein proteins, and and their influence activators, interacting proteins, can transcriptional HSFB2B finger not and they are zinc HSFB1 HSFs that stress-associated B facil- HSPs, suggest class factors as While interactions HSFA1-type such NPR1. to genes with bound interaction target it through of where number monomers, pathogens. a (Olate plant in HSFA2 of to nucleus response induction the SA-mediated the entered of itating regulator NPR1 key temperatures is which low NPR1, In with reported was factors HSFA1 05 Perez-Salamo 2005, , tal. et 08.Teedt eeldta SA n SA a rmt odacclimation cold promote can HSFA2 and HSFA1 that revealed data These 2018). , tal. et tal. et 07.Rcnl SAAadHF5wr eotdt oto together control to reported were HSFA5 and HSFA4A Recently 2007). , 04.Erirw eotditrcino SAAadphosphorylation and HSFA4A of interaction reported we Earlier 2014). , tal. et 00) h D0rpa OLS TL n TPL-related and (TPL) TOPLESS repeat WD40 The 2010b). , tal. et 15 tal. et 09 Perez-Salamo 2019, , 09.Itrcino A7wt HSFA4A with BAG7 of Interaction 2009). , BAG6 tal. et a dnie among identified was 04.T identify To 2014). , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. etsoktasrpinfco novdi eeomna euaindrn mroeei nsunflower. in embryogenesis seed-specific during A (2002) regulation J. developmental Jordano in & Chem R. involved Biol Carranco P., factor Prieto-Dapena transcription J., Diaz-Martin heat-shock A., Rojas C., seedlings Almoguera in J. stress Jordano oxidative & and dehydration A. tobacco. extreme Diaz-Espejo transgenic from M., apparatus of Lindahl photosynthetic The the J., Tejedor-Cano of (2009) functional Protection J.M., (2012) Personat J. through P., seeds Jordano Prieto-Dapena C., of & Almoguera longevity R. and Carranco thermotolerance J.M., HaHSFA9. basal with Espinosa enhances interaction J., factor Diaz-Martin transcription P., HaDREB2 Expressed Greening. Factors Prieto-Dapena Seedling Stress and C., Heat Longevity (2020) Almoguera J. Seed Jordano Regulate & Differentially development J.L. Maturation stress, Ruiz Seed R., cell during Carranco of P., integrators Prieto-Dapena factors: C., Almoguera shock Heat and (2010) heat L. to Sistonen lifespan. response & and in R.I. wheat Morimoto from M., Akerfelt HSFs of characterization conditions. Functional stress (2019) abiotic P. other Khurana & P. interacting Agarwal regulation, in differences REFERENCES appropriate their 6. the with and networks Other genes. interactions similar stimuli. target regulatory biotic in and and illustrates proteins positioned and abiotic 5 various genes are to Figure target HSFs responses of stress. plant coordinates methylation regula- which to histone HSFA4A, Epigenetic promotes response of network which elevated HSFA2, transcriptional proteins. of by such defense-related capability controlled proteins or is the transcrip- protective memory maintains other regulators stress encode include metabolic heat can can scavengers, of which genes ROS tion genes target target chaperons, Other of genes. forms, HSPs, cascades. range target trimer as a transcriptional of in of forming transcription transcription motifs regulators, activate induce tional HSE HSFs to regulatory capacity trimers. HSF their cis active influence to or can HSFA2 bind multimers as HSFs H including HSF such interactions of HSFs formation. stability protein-protein of H trimer therefore or subset particular heteromeric sumoylation of in and ROS, subsequent phosphorylation responsible homomeric partially influence in conditions. least resulting can adverse at cascades, Phosphorylation in are HSFA4A. kinase signals genes ROS MAP HSF way. re- stimulate stress-induced multiple transcriptional signals in of and signals activation hormonal ROS transcriptional metabolic, integrate sensitive for induce to a can seem is accumulation which HSFs ROS homeostasis, ox- sponses. oxidases. changing photosynthetic generates NAPD of in membrane-bound infection indicator of breakdown and Pathogen activity metabolic accumulation to the ROS transports. due triggering to by electron ROS particular burst stresses mitochondrial generate idative biotic in and stresses and homeostasis, environmental photosynthetic abiotic cellular All disrupted other in activity, of changes balance. number to redox a fast in to very only alterations responses respond not control can large are but HSFs and HSFs tolerance, and plant regulation also. stress that biotic transcriptional heat demonstrate, to post conditions, evidence regulators and environmental Accumulating key whose transcriptional changing interactions. system on regulatory protein-protein to reflected multilevel of is complex, responses spectrum variability a molecular of HSF part coordinate stresses. is to abiotic plants in is factors functions shock heat principal the HSFA4A. of with world association diverse The their to function precise characterization CONCLUSIONS Further assign HSFA4A. to 5. with and associated interactions interacting been validate HSFA4A attacks previously to that pathogen have needed and show, which stress is results death, to Our responses cell homeostasis, programmed unknown. cellular and is control, transcriptional function in its involved are but proteins transduction, signal in implicated 2 O 2 a xdz y mn cdrsde,pooefraino y-y od n hrfr tblz the stabilize therefore and bonds Cys-Cys of formation promote residues, acid amino Cys oxidize can , 277 a e o elBiol Cell Mol Rev Nat 43866-43872. , LSONE PLoS M ln Biol Plant BMC uc nerGenomics Integr Funct , 7 , 11 e51443. , 545-555. , , 9 75. , 16 , 19 497-513. , lns(Basel) Plants , 2 9 O . 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Sumoylation acquired (2010) during A. activity Avni its & modifies A. (HsfA2) Breiman A2 D., byfactor Meiri S., phosphorylation Schuster Sequential heat R., by (1996) Cohen-Peer activation S.K. transcriptional represses Calderwood 3 & kinase factor-1. M.A. synthase shock Stevenson glycogen and B.D., kinase Price protein and mitogen-activated F., stress abiotic Soncin species, B., oxygen Chu Reactive (2017) R. Mittler & combination. E. stress Blumwald R.M., Rivero evolution F.K., the shaping Choudhury regulator interactions: a response. Host-microbe immune ICE1: (2006) B.J. plant (2003) Staskawicz the J.K. & B. of Zhu Day & G., Arabidopsis. Coaker M. S.T., Agarwal in Chisholm X., tolerance Hong freezing B.H., and Lee transcriptome reveals S., cold-induced Kanrar profiling of Transcriptional M., Ohta (2002) V., Arabidopsis. S. Chinnusamy in Luan responses & hormonal T. and and Zhu stress, Identification abiotic X., Physiol Functional pathogen, Wang wounding, R., (2016) between Cadmium Gupta B. interactions and novel H.S., Huang Salinity Chang & Conferring Y.H., Z. Paspalum Cheong Yang Seashore L., Halophyte Zhuang from J., Cloned Liu Tolerance. Genes Z., Tan of C., SaHsfA4c Characterization (2020) Chen R. Zhuo Y., & X. Chen Han Heat L., and Wu Activities G., ROS-Scavenger Qiao Expression. Regulating M., by Proteins Liu Tolerance Y., Shock Cadmium Zhang Enhances Z., Hance Lu alfredii Y., Sedum Wang as From H., functions Li M., DREB2C Yu Arabidopsis S., Chen (2010) response. C.O. stress Lim heat & the S.Y. during Lee HsfA3 238-244. D.Y., of Kim activator C.J., transcriptional Lim a J.E., Hwang H., Chen hua . hrn . gra .&KuaaP 21)Ha hc atr nrc Oyastv L.): sativa (Oryza rice in factors shock Heat stress. abiotic (2011) and P. development Khurana reproductive , & during Arabidopsis. P. analysis Agarwal in expression N., genome-wide thermotolerance heat-inducible Khurana acquired A H., (2007) of Chauhan T.T. extension Wang & for S.H. required Chang C.N., is Wang Physiol The W.T., HsfA2, (2018) Chi factor, S. N.Y., Repair. Liu Amiard transcription Damage H.C., & Liu DNA F. Y.Y., and Butter Charng Stability C.I., Telomere White in F., Involved Benyahya Is , O., GH1-HMGA1 Ines of Histone Da activation Linker O., synergistic Rymarenko for C., complexes Charbonnel superactivator between interaction expression. hetero-oligomeric Specific gene gene (2009) creates stress for K.D. HsfA2 Scharf heat framework & and a L. Nover HsfA1 interactome: D., tomato TOPLESS Bublak S.K., The Baniwal (2012) K.Y., Chan-Schaminet B. Davies & Arabidopsis. W. in Guo repression M., Ashworth B., Tolerance. Causier Desiccation Involving and Synergism Longevity SUMO-Dependent Seed to (2017) controls Sci Linked J. that Functions Plant Jordano gene with Factors & NPR1 Transcription C. Arabidopsis Shock Almoguera Heat The repeats. P., ankyrin (1997) Prieto-Dapena containing X. R., protein Dong Carranco novel & a S. encodes Volko resistance J.D., acquired Clarke systemic J., Glazebrook H., Cao 177 286 311-327. , 171-187. , , , 129 143 , rn ln Sci Plant Front 8 974. , 661-677. , 251-262. , ilChem Biol J ln J Plant , ln Physiol Plant 7 rn ln Sci Plant Front ilChem Biol J , , 90 102. , Cell 271 856-867. , , 30847-30857. , 124 , 803-814. , , 158 284 , 11 423-438. , 20848-20857. , 142. , 18 ice ipy e Commun Res Biophys Biochem ee Dev Genes ln o Biol Mol Plant , Cell 17 1043-1054. , o ee Genomics Genet Mol , 88 , 57-63. , 74 ln Physiol Plant 33-45. , , Front Plant Plant 401 , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. aaoD,Ss . akiI,AdaiN zbdsL 21)PatieOesa ffral,Non- Arabidopsis. Affordable, in thermotolerance an (2018) acquired M.M. Offers long-term Castellano in PlantSize & role H.P. (2018) major Mock 41 a R., L. plays Toribio Szabados family A., HOP Vitro. Munoz & in L., Fernandez-Calvino N. Color N., and Andrasi Fernandez-Bautista Size Plant I., Measure Valkai to L., Method destructive Sass D., Farago D. Wagner & Y. Eshed J.C., responses. Hong cytokinin K.D., of Birnbaum modulation E., chromatin-mediated Steiner by M.F., maturation Wu , leaf Shock H.J., of Heat Kim Regulation Between S.K., (2013) Heat. Interactions Han to Multi-Level I., Acclimation (2015) Efroni Regulate I. System Rieu Redox & the S. and Identification Park 999. Proteins, (2006) J.L., Shock M.B. Peters Heat Dickman the thaliana. J., Factors, & in Xu Arabidopsis J. motifs N., in Reed AHA Driedonks family A., of protein Godzik role BAG E., The the Zalm (2000) of 18793-18801. der L. characterization van Nover functional HsfA2. S., & and and Chen A. HsfA1 factors E.V., Chen transcription Doukhanina R., stress Lyck heat Proteome- tomato C., (2019) A., of Kistner Pasha N.J. function Provart activator A., E., Treuter Gaudinier & P., H., B. Nahal Doring Usadel T., R., Viewer. Sivieng Grene Interactions Y., Molecular S.M., Wu Interactions/New Brady Physiol Protein-Protein E., of M., Esteban Prediction Tyers M., Structure-Based K., wide, Ierullo Dolinski R., R., Song Oughtred V., ROS Lau of photosynthetic NineTeen S., with Importance the Dong metabolism (2010) of mitochondrial K. Recruitment of Padmasree (2016) interactions & X. beneficial assimilation. A.S. Cao the carbon Raghavendra & during S.R., C. system antioxidant Yearla Liu and V., X., Abhaypratap Kong C., J., AtPRMT5. Dinakar Sun requires L., spliceosome activated Gu the L., to Wang Complex T., Lu X., Deng vadA,KmrM,Lcuiu . uk . o okl-oigP itH 21)Rglto fthe of Regulation (2013) HsfA2. factor H. stress Hirt heat & the P. of Koskull-Doring phosphorylation MPK6-targeted von by J., , Arabidopsis Lucks in D., response from stress Lecourieux heat (HSFs) M., factors Kumar transcription Functional A., shock (2019) heat Evrard by S. recognition Fragkostefanakis DNA in properties. & Diversity organisms. domain (2011) E. model binding H. Schleiff Sakurai DNA & on K.D., Y. based Enoki Scharf is factors S., HsfA1 Simm tomato of S., diversification Ullrich A., El-Shershaby 1. factor shock heat of mammalian network the gene of oxygen Regulation reactive (2017) A.T. the J Dinkova-Kostova & of S. Mittler component & Naidu central Dayalan K. Schlauch a V., is Shulaev 1 J., Coutu peroxidase D.J., Arabidopsis. ascorbate Oliver and Cytosolic Z., transcription Shengqiang (2005) inhibit H., R. 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No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. itre nisros ih-addsatn,adgairps ensagn htecdsaheat-shock Arabidopsis a of encodes mutant that new gene A a (2008) defines F. gravitropism Migliaccio and & slanting, S. right-handed Ferrari roots, factor. P., its Tassone in S., disturbed Piconese A., Fortunati uZQ ogX 21)Ssei curdrssac:triglclifcinit lbldefense. global into infection local turning resistance: acquired paralogs Systemic Biol protein (2013) Plant factor-binding X. Rev factors. shock Dong shock heat & heat maize Z.Q. specific The Fu with (2006) non-redundantly M.J. interact Scanlon HSBP2 & and L. EMP2 Nover P., meta-analysis. Rogowsky microarray S., com- and Fu network profiling Chaperone transcriptome (2015) E. by Schleiff Environ explored & K.D. Cell lycopersicum Scharf Solanum D., in Bublak P., position Paul S., Simm S., Fragkostefanakis erlt . oigP,BneisF,Wnehu .&NvrL 20)Teblneo ula import nuclear of factor balance transcription The stress (2001) heat L. tomato of Nover transcript function & nodule-specific and S. the distribution Winkelhaus intracellular of HsfA2. the F., Maturation determines Bonzelius (2008) export P., G.Q. and Doring Yu & D., trans-splicing. interallelic J.B. Heerklotz involve a Zhu may of sativa Y.Z., splicing Medicago Wang alternative in H.S., and MsHSF1c Zou Structure (2007) Z.S., G.Q. He sativa. Yu Medicago & in J.B. MsHSF1, Zhu gene, Y.Z., 1056-1061. factor Wang transcription H.S., shock Zou element. heat R., shock Xie heat Z.S., the He implicated of is architecture factor transcription the shock on heat 3648-3659. yeast dependent and the activation chaperones of Hsp70 gene-specific Phosphorylation and (2004) in H. Hsp90 Sakurai between & (Capsicum Crosstalk N. Hashikawa (2011) tomato. pepper K.D. in in Scharf factors expression & CaHsfA2 transcription analysis, E. stress of Schleiff Genome-wide heat characterisation D., (2015) Bublak and M.H. A., (CaHsfs) Lu Hahn family & Z.H. gene Transcription Gong factor Stress W.G., L.). shock Heat annuum Chai heat Plant Y.F., The of Zhai (2016) profile J.P., M.H. Lu Stresses. Lu M., Abiotic & Guo to Z.H. Response Gong in Function D.X., and Luo 114. Regulation, X., Structure, Ma (HSFs): J.H., Factors Liu Arabidopsis. in M., Thermotolerance Guo and Part- Interacting (2014) Regulation Its Gene M.F. and RCF2 Stress-Responsive Yi Phosphatase Cell Heat Protein & LlHSFA2 for Plant The J.N. Critical (2014) with J.H. He Are interact Zhu NAC019 S.S., & can ner H.T. Seng Zeng longiflorum), X.L., X.H., thaliana. Yue (Lilium Zhong Q.M., Arabidopsis lily Guan Z., transgenic in Wu of factor thermotolerance M.A., the transcription Khan enhance stress J.J., and and heat Sui factors novel J., transcription Wu a shock LlHSFA1, J., heat Yi of B.H., Regulation Gong disease. (2018) and D.J. physiology Thiele in & roles their E.T. rhodopensis. Burchfiel V. Haberlea Toneva R., relic & Gomez-Pastor glacial A.R. A.S., resurrection Fernie Sci Marriott the J.H., M.R., Life in Temanni Schippers Mol tolerance J., B., desiccation Cell Hille Mueller-Roeber of I., mechanisms Minkov C., Molecular N., Antonio (2013) Sujeeth J., T., Thomas-Oates Tohge reveals E., T., kinases Obata Bergstrom protein M., acid-activated Benina abscisic T.S., stress. 3 Gechev and in deficient reproduction, mutant growth, Arabidopsis in (2009) roles J.K. critical Zhu & H. Fujii x Bot Exp J o elBiol Cell Mol , 26 M ln Biol Plant BMC , 38 , 438-453. , 64 , 693-709. , , 70 59 839-863. , 689-709. , , 1363-1374. , 21 1759-1768. , , 15 151. , a e o elBiol Cell Mol Rev Nat ln Cell Plant rcNt cdSiUSA S U Sci Acad Natl Proc 20 , 23 741-755. , , 19 4-19. , Genomics ice ipy e Commun Res Biophys Biochem ln elReports Cell Plant , 106 Planta , 92 8380-8385. , 115-121. , , o elBiol Cell Mol 224 rn ln Sci Plant Front , 33 42-52. , 1519-1533. , , , Annu 364 Plant 24 , 7 , , , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. .&Fakseaai .(00 aua aito nHf2pemN piigi soitdwt changes with associated is seed splicing Schleiff for K.D., pre-mRNA required Scharf HsfA2 A., domestication. is in Bovy tomato variation D., and during Natural Bublak thermotolerance nucleus P., (2020) in Gebhardt the S. S., Fragkostefanakis Roth to & J.M., translocation E. Jimenez-Gomez A., by Mesihovic Y., response Hu shock heat functions of AtHSBP, Arabidopsis. protein, 230 in regulator factor-binding development serine shock negative heat of Cytosol-localized a (2010) Phosphorylation T.L. as (2001) Jinn Mor- & 1. L. H.C. J., factor Lai Sistonen Hellman S.F., shock & Hsu A., heat J.E. Meinander of Eriksson activity M., R.I., transcriptional Kallio inducible Morimoto J.O., promotes C., Rantanen MacKintosh A., N., Mikhailov rice V., Hietakangas C.I., Holmberg in . hn,Q,Ce,L,Lag . u .(08 coi vrxrsino az etsoktran- shock heat Arabidopsis. maize transgenic of in overexpression tolerance Ectopic salt-stress (2018) and J. thermo Wu, Plantarum increased Physiologiae Y., confers Acta LIang, ZmHsf04 factor L., gene transcription factor Chen, shock scription Q., heat Zheng, Drosophila Y., of Jiang functions Multiple (1997) C. Wu vivo. & in Takahashi M.A. Reprogramming Cellular D.S., Mortin Favero for P., Network D., Jedlicka Regulatory Coleman Gene A L., (2018) Watt K. Regeneration. A.M., Sugimoto Plant Bagman & in A., S.M. the Brady Iwase of S.E., B., Ahnert repressors Rymen T., as M., Shibata act M., thermotolerance. HsfB2b Ikeuchi acquired and the HsfB1 regulate Arabidopsis I positively 1243-1254. (2011) photosystem but , M. Hsfs photosynthesis, Ohme-Takagi heat-inducible on (2004) & of D. effects N. expression Leister Mitsuda functions. thaliana: & chloroplast M., F. for Arabidopsis Ikeda Salamini genes of P., nuclear D Jahns of expression A., subunit and Schneider I stability in E., photosystem photosystem Richly Lhca4 Impaired for C., protein (2008) Varotto Mutants P., I D. Pesaresi complex Leister A., light-harvesting & Ihnatowicz the B. of Muller phosphorylation D., thaliana. Wolters STN7-dependent Arabidopsis K., induces Lohrig oxidation P., I Pesaresi A., Ihnatowicz stress. abiotic factors to shock Heat response (2015) A.S. profiles Xiong expression & J. and Ma classification, G.L., Rep Wang identification, W., Biol genome-wide Huang Z.S., carrot: Xu F., in hit2/xpo1a Wang AtHsfA4a. M.Y., Arabidopsis of Li of confinement Y., phenotype nuclear Huang sensitive by irradiance part High in (2018) caused S-J. is Wu, mutant K-Y., Chang, H.-Y., Huang identified protein heat-stress- a regulating HsfA1d, by (2013) tolerance T. heat Taji improves & Ishida cDNAs T. K., expression. salsuginea Hayashi Yamaguchi-Shinozaki gene Thellungiella Y., K., responsive Sakata using Kusakabe K., hunting A., Shinozaki FOX Shimura M., via T., Seki Katori M., T., Tanaka Ishikawa J., EDS1 N., Arabidopsis Ohama (2011) Y., J.E. Higashi Parker responses. & immune L. compartment-specific Deslandes cell to C., 1404. recognition Pouzet effector C., pathogen Tasset connects L., Wirthmueller K., Heidrich wn .. i .. o .. en .. o .. ktrS,Co ..&Bh ..(2014) J.D. Bahk & G.J. Choi S., salinity high Akhter under Y.S., factor Son transcription ABA heat-shock Connects H.S., a HSFA6b conditions. Jeong as Factor dehydration HsfA6a M.S., and Stress Arabidopsis Woo Heat of D.W., The characterization Functional (2016) Kim T.L. S.M., Jinn Hwang & Responses. C.R. Heat Yang ABA-Mediated and C.Y., Signaling Niu Y.C., Huang MOJ EMBO , 42 893-905. , , 16 Planta 2452-2462. , ln elPhysiol Cell Plant o Plant Mol ln elEnviron Cell Plant ln Physiol Plant , , 227 40 9. , 717-722. , , 6 411-422. , . , 153 ln Physiol Plant , 773-784. , 37 e Phytol New 21 1202-1222. , , 172 , 225 ilgaPlantarum Biologia MOJ EMBO ln J Plant 1182-1199. , 1297-1310. , , 37 , 20 839-852. , 3800-3810. , , 62 Science ln Physiol Plant 69-79. , , 334 1401- , , 157 Mol Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. e .. edro ..&ZuJK 20)TeAaiosscl-epnietasrpoeadisregu- its and transcriptome cold-responsive Arabidopsis The (2005) J.K. Os- ICE1. Zhu by & & lation C.M. D.A. Henderson Malmstrom to B.H., F., compartment. help Lee Brandizzi chloroplast thaliana the Arabidopsis C.A., of from Sinkler size genes A.K., COVERAGE the CHLOROPLAST Stavoe establish REDUCED M.E., (2016) Ruckle K.W. shock G., teryoung seeds. heat Stefano a in as R.M., tolerance BnHSFA4a Larkin of desiccation characterization of Functional (2017) re-establishment X. the Wang sustained & controlling 2361-2375. governs X. in factor Li factor shock H., Xue heat transcription X., hit-and-run Liu A S., (2016) Lang memory. I. stress Baurle transcriptional & between and S. methylation Cross-talk Altmann histone (2007) K., Brzezinka K. J., Apel thaliana. Lamke Arabidopsis A & in S I. responses U stress Murgia Sci of E., Acad signaling Warzych Natl peroxide-dependent hydrogen E., and Pers-Kamczyc oxygen- M., singlet comprehensive Stachowiak reveals light. C., Metabolomics UV-B (2011) Laloi to A.R. Fernie Arabidopsis & of K. responses 354-369. Saito Kawashima metabolic , H., M., independent Goto two Matsui Y., involving Kondou R., H., reprogramming Niida Otsuki N., F., Hayashi Arabidopsis. Matsuda M., Kobayashi in M., A., resistance Fukushima T., pathogen Tohge HsfB1 and M., factors Kusano expression shock Pdf1.2 Heat of (2009) F. regulation Schoffl the & in Plant T. involved Nurnberger are B., Kemmerling HsfB2b H., and Birke W., Busch M., Kumar to thaliana Arabidopsis of Responses (2008) S.Y. Lee & D. Mackey W.Y., syringae. Kim Pseudomonas by S.Y., challenge Arabidopsis Kim in factor M.G., splicing stress-specific Kim heat a as STABILIZED1 kDa (2018) Y.H. 70 Cho and thaliana. & 1 of S.D. factor Yoo inhibitor transcription G.D., an Kim shock 1, heat factor Arabidopsis shock between proteins. Interaction Heat shock (2002) (2015) heat F. Y.S. Schoffl Lee & & B.H. (2013) H.J. Kim Cha J. N.Y., Chory repair. Gil joining & H.J., end B.J. Lee light. non-homologous excess E.H., Pogson to Kim T.C., Arabidopsis G.Y., of Mockler Kang A response family F., S early factor Hong the U transcription for Sci B., Acad required shock Cole factors Natl heat transcription G.M., rice heat-shock Estavillo of of P.A., Subset view Crisp systematic H.S., A Jung (2013) K.H. Jung analysis. & phylogenomic H.J. using Gho G.H., Jin xrsino etsrs rtisdrn eddvlpeto Arabidopsis. of development regulating seed cascade transcriptional intracellular during novel and proteins A (2007) stress function P. Koskull-Doring heat activator von of for & com- H. expression essential Baumlein signature motifs E., Vierling new NES C-terminal S., a Kotak and of of AHA Characterization identification with (2004) and Hsfs P. (Hsfs) A Koskull-Doring localization. factors class von transcription plant & of stress F. bination heat Bicker Arabidopsis A., directs of Ganguli PRR7 domains M., of repression Port HsfB2b-mediated S., clock. (2014) Kotak circadian S.A. the Kay of & responses to J.L. stress Responses Pruneda-Paz Network.abiotic Rapid B.Y., Transduction (2019) Chow Signal R. E., the Mittler for Kolmos & J. Landscape Kangasjarvi the M., Priming Nuhkat Stress: S., Sengupta Abiotic S.I., Zandalinas H., Kollist , 2 ln inlBehav Signal Plant 152-165. , ln J Plant ln Cell Plant x Bot Exp J , , , 39 104 110 98-112. , , 672-677. , 14474-14479. , 17 ln Physiol Plant J , 13 , 3155-3175. , 53 e1432955. , Oncotarget 371-375. , o Cells Mol , rcNt cdSiUSA S U Sci Acad Natl Proc , 170 , 6 25 29712-29724. , rcNt cdSiUSA S U Sci Acad Natl Proc 321-329. , 323-331. , 22 MOJ EMBO , 35 162-175. , , 111 ln Cell Plant , rnsPatSci Plant Trends 113 16172-16177. , E1116-1125. , , 19 x Bot Exp J 182-195. , ln J Plant , 24 25-37. , , Proc Proc 68 , Mol 67 , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. iM,Dl . ekranK,OcigC,BrnznKW co .(00)Dtcino nvivo in novel of of Detection identification (2010b) and F. fragment, Schoffl BiFC & novel K.W. a Berendzen proteins. using interacting C., A-HSFs, B-HSF late Oecking class class and K., Arabidopsis early Weckermann between between J., interactions interactions Doll and specificity M., Promoter Li (2010a) F. factors. Schoffl shock & heat Arabidopsis K.W. Berendzen of tolerance M., maize drought-stress of Li and Expression (2015) thermotolerance X-L. Sheteiwy the Guo, enhances & H-M., ZmHsf06 Arabidopsis. Zhang, X.L. gene Y-M., transgenic Zhang, Guo factor Z-H., transcription in L.N., Liu, shock thermotolerance G-L., Zhao heat mutant. improves Li, athsfa2 L. H-N., Y.J., mays Zhang, the Zhang H.-C., Zea of Li from defects Y.Y., factor thermotolerance Zhang transcription rescues shock G.Y., and heat Wang thaliana new Arabidopsis H., a ZmHsf05, Shao (2019) CmHSFA4 H.N., Chrysanthemum M.S. (2018) Zhang S. G.L., Chen & Li chrysanthemum. F. Chen transgenic J., in Jiang tolerance A., stress Song 1321. salt T., regulates Gao positively H., Zhao gene H., Zhang F., factor Arabidopsis. Li shock transgenic heat in engineered thermotolerance genetically increased of and activity J proteins the shock Plant of heat Derepression of (1995) synthesis F. constitutive Schoffl causes & A. Hubel J.H., Lee iaaL rhr .(06 h Itc rjc n oeua neato Databases. Interaction Molecular and Project MIntAct The (2016) S. Orchard Biol & transcription transgenic L. stress in heat Licata germination a SlHsfA3, in of hypersensitivity overexpression salt Ectopic and (2013) thermotolerance J. Arabidopsis. increased Li confers & tomato, X. Xu from A., factor 7 Wang L., athanogene Zhang (Bcl-2)-associated Z., Li lymphoma2 WRKY29. to B-cell binding Arabidopsis and (2017) sumoylation M. translocation, , pathway. requires Dickman tolerance regulation & heat heat B. 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Liu Zhang Roitsch the & & J. of Hu R. regulator B., Ehness possible Liu R.K., a Proels and tomato: M.G., HSFA4a, in HSFA2, Hofmann 179-183. Arabidopsis P., kinase of Vashista MAP roles A.K., activated The response. Sinha protein (2018) cytosolic V., C.H. Yeh and Link & response S.J. shock Wu heat C.A., the Lu in HSFA7a M.Y., Tsai K.F., Lin 214 , 95 , 695-705. , 1415 401-413. , , M Genomics BMC 8 , 603-612. , 55-69. , 39 LSONE PLoS 39-67. , ucinlPatBiology Plant Functional , , 8 15 e54880. , 1009. , ln o Biol Mol Plant u elBiol Cell J Eur , 73 , , 42 559-567. , 89 23 1080-1091. , 126-132. , ln Physiol Plant o Stud Bot , 163 ln itcnlJ Biotechnol Plant , 59 276-290. , ln Sci Plant 15. , ESLett FEBS caPhysiologiae Acta , 283 ehd Mol Methods e Phytol New , Cells 16 375-384. , , 1311- , 531 Plant , 8 . , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. aaauM,RdyPS,KmrSA,Siatv .. ihrPBK a ..(05 Genome-wide (2015) D.M. Factors. Rao Transcription Shock & Heat P.B.K. L. Kishor bicolor R.K., Sorghum , Srivastava of acclimation. S.A., Characterization acquired Kumar and systemic P.S., Scanning in Reddy ABA M., and ROS Nagaraju of roles The L.). sativa (2015) E. (Oryza , Blumwald rice & in interactions R. and factors Mittler affinities shock Binding heat (2011) and A. 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Data may be preliminary. oec .. oec .. ahnS,ZagZ,Grti . ndrM iehKmrSP (2009) S.P. Dinesh-Kumar microarrays. & protein M. functional Snyder using M., revealed Gerstein thaliana (1999) Z., Arabidopsis H.C. Zhang , in Nelson S., networks Bachan target & G.V., MAPK D.S. Popescu transcription King shock S.C., heat D.E., Popescu the Wemmer from C.W., domain trimerization Liu the Y.K., of factor. characterization Shin biophysical to and tolerance M., Biochemical seedling Rabenstein on effects R., synergistic Peteranderl in of and Co-overexpression stress. longevity oxidative (2014) seed J. and enhanced dehydration Jordano in severe & results Factors C. causes Shock Almoguera MYB44 P., Heat factor two Prieto-Dapena transcription J., Tejedor-Cano Arabidopsis J.M., stress. by Personat A4A abiotic repression to factor Dominant hypersensitivity shock (2014) and MPK3 heat A. damage kinases oxidative The Pitzschke protein Domoki & (2014) mitogen-activated B., H. L. the Horvath Persak and Szabados I., stress & oxidative Nagy C. by V., Koncz Transcription regulated Lumbreras MPK6. is L., Shock and B., and Bogre Heat Vilela tolerance Novel K., L., salt A Medzihradszky confers Zsigmond (2013) Z., G., Darula Rigo y. Stresses. M., C., Wang Abiotic Papdi and M., I., Biotic He, Perez-Salamo in Y., Involved Yang, is pseudoreticulata Report J., Vitis Biol Shi, Wild Mol K., Chinese from Zhao, VpHsf1, Z., G., update. Factor, K. Leung 2019 Dolinski Z., L., A., database: S. O’Donnell Chatr-Aryamontri interaction Peng N., J., BioGRID Kolas Coulombe-Huntington The C., A., (2019) of Chang Willems M. release L., S., Tyers Boucher the Dolma & J., F., after Rust Zhang responses Gobel B.J., R., E., stress Breitkreutz McAdam Hideg distinct C., D., of Stark Wagner induction R., A., Rapid Oughtred Danon (2003) C., K. Arabidopsis. Kim Apel in C., & oxygen Laloi M. singlet C., Nater Ochsenbein I., D., pathway Feussner Przybyla regulatory C., novel R.G., a Camp mediates factors. den HSFA1 NPR1 op with (2018) interacting J. by Salinas & acclimation L. cold Holuigue in J.M., of Jimenez-Gomez Network E., Regulatory Transcriptional Olate (2017) K. Yamaguchi-Shinozaki Response. & Stress K. Heat Shinozaki Plant H., of Expression. Factor Sato Response Transcription Stress N., of Heat Ohama Level S., Yanagisawa the the T., in at Ishida Cascade Regulated F., Transcriptional Takahashi Strictly The S., Is (2016) Koizumi Arabidopsis S., K. Yamaguchi-Shinozaki Kidokoro & H., Zhao K. J., Shinozaki gene Mizoi K., HsfA2 Kusakabe Arabidopsis N., growth. Ohama the callus enhanced of and overexpression tolerance stress High-level salt/osmotic Bot also (2007) Exp but T. themotolerance increased Nishiuchi only & not confers K. Yamaguchi D., The Ogawa (2006) K. Arabidopsis. of Apel responses & stress I. Feussner oxygen-mediated J singlet A., during Imboden Plant susceptibility) C., disease Gobel (enhanced world: F., EDS1 Hsf Landgraf of role The A., Danon factors. (1996) D., transcription W.B. Przybyla C., Gurley stress Ochsenbein & heat plant E. Czarnecka-Verner of P., genes Vergne properties target D., and of Gagliardi classification regulation K.D., the Scharf HsfA2. of L., factor, Analysis Nover transcription (2009) shock S. heat Shigeoka Arabidopsis & an Y. by Yabuta E., Yoshida A., Nishizawa-Yokoi otM,TipJ,ZeisiD,WbrC,HekozD,Wnehu . ulkD cafKD (2004) K.D. Scharf & D. factor Bublak transcription stress S., heat Winkelhaus tomato of D., factor Heerklotz retention cytoplasmic C., and HsfA2. Weber coregulator as D., Hsp17.4-CII Zielinski of Role J., Tripp M., Port 23 80-92. , Biochemistry ln Physiol Plant , , 47 58 ln Physiol Plant 445-456. , 3373-3383. , , 31 , 38 240–247. , , 135 3559-3569. , , 165 rnsPatSci Plant Trends 1457-1470. , ln Cell Plant 319-334. , M ln Biol Plant BMC , 15 , 2320-2332. , 22 53-65. , 25 a Plants Nat n o Sci Mol J Int , 14 isiBoeho Biochem Biotechnol Biosci 56. , , 4 uli cd Res Acids Nucleic elSrs Chaperones Stress Cell 811-823. , , 15 2517-2537. , ln Cell Plant , 73 , 47 890-895. , , D529-D541. , , 28 1 ee Dev Genes 215-223. , 181-201. , Plant J Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. cafKD,BreihT,EesegrI oe .(02 h ln etsrs rncito atr(Hsf) factor transcription stress heat plant The (2012) evolution. L. and Nover shock function & heat structure, I. the family: Ebersberger of T., regulation Berberich Negative K.D., (1998) Scharf R.I. Morimoto & HSBP1. J.M. by modulation Kramer response chromatin S.G., transcriptional Fox recognition, D., DNA Chen factors: S.H., shock Satyal heat of aspects Novel expression. (2010) gene Y. and Enoki gene & heat-stress-responsive H. and Sakurai water-stress-responsive are in function function DREB2A Dual (2006) factor repressor K. Yamaguchi-Shinozaki expression. transcription and & K. Arabidopsis DNA-binding Shinozaki HsfB1. Y., an (2017) factor Osakabe of F., transcription E. Qin stress Schleiff K., heat Maruyama & (2014) Y., tomato Sakuma Y. K.D. the Scharf Saijo of turnover & D., the K. Bublak for Tsuda O., prerequisites immunity. Y., Mirus plant Takano systemic S., X., and Roth Lu local M., couples Yamashita-Yamada pathway cytosolic K., PEPR for Arabidopsis Hiruma required The K., is Zat12 Arabidopsis. Yamada protein in A., finger stress Ross zinc oxidative The during nucleolar (2004) expression R. and 1 Mittler response & peroxidase H. stress ascorbate Liang between S., Davletova link L., new Rizhsky protein. A AtREN1 by (2014) mediated D. Arabidopsis Honys in 670-683. & development K. pollen kinase Solcova during vitro. cyclin-dependent function A., in a Gibalova HSF1 by D., factor Phosphorylation Renak transcription (1997) L. heat-shock Bako Arabidopsis & the C. of 115 Koncz binding J., factor DNA Schell transcription modulates shock F., Schoffl heat thaliana A., Arabidopsis Reindl TBP. the protein in between binding TATA HvHsfB2c Interaction the factor (1998) and shock F. development. HSF1 heat Schoffl seed the & and Sreeniva- by A. response & proteins Reindl M.K. stress shock Reddy drought heat J., in small Lee implications the L., its of Eschen-Lippold regulation barley: M., Unraveling Innate Kuhlmann (2014) Arabidopsis C., N. in Seiler sulu P.B., Cascades Kishor Kinase Kavi MAP P.S., (2012) Reddy J. Mundy & M. Petersen Immunity. M., expression. HsfA1a Roux Arabidopsis Hsp M.W., of Rasmussen Overexpression stress-induced heat (2014) promoting and L.M. Zhang oxygen by & low W.P. tolerance Research Li in stress L.L., element Yang diverse Y.F., common Liu enhances as J., signaling Chen J., ROS Qian (2012) P. seed- Perata a & of V. stresses. overexpression Banti ectopic C., organs. The vegetative Pucciariello in (2008) dehydration J. severe to Jordano tolerance & , confers HaHSFA9, C. factor, Almoguera transcription deteri- R., specific controlled to Castano resistance P., Improved Prieto-Dapena (2006) J. Jordano seeds. & transgenic C. in Almoguera oration R., Castano transcription photomorphogenesis. P., Seed-specific early Prieto-Dapena (2017) J. and Jordano embryogenesis & late F. links Merchan HSFA9 J.M., from factor Personat factor overexpressed C., shock Almoguera when P., heat thermotolerance Prieto-Dapena new confers a and HSF3, response plants. (1998) shock transgenic F. heat in Schoffl the & derepresses G. thaliana, Eggers-Schumacher Arabidopsis K., Hinderhofer R., Prandl 54 93-100. , 1004-1014. , ln hso Biochem Physiol Plant , 13 rn ln Sci Plant Front rcNt cdSiUSA S U Sci Acad Natl Proc 1233-1243. , o e Genet Gen Mol ESJ FEBS , ln Physiol Plant 3 169. , , 277 , 59 ee Dev Genes 4140-4149. , 3-10. , , , 258 103 ici ipy Acta Biophys Biochim , 269-278. , ESLett FEBS 142 18822-18827. , , 1102-1112. , 12 1962-1974. , 26 , 436 318-322. , , 1819 x Bot Exp J 104-119. , ilChem Biol J LSONE PLoS MOJ EMBO , ln elEnviron Cell Plant eeisadMolecular and Genetics ln J Plant 68 , 1097-1108. , , , 9 279 33 e89125. , ln Physiol Plant , 62-75. , 11736-11743. , 89 31-44. , ln J Plant , 37 , , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. ogrPK ehmHR 18)Yatha hc atri nesnilDAbnigpoenta exhibits that Cd- protein DNA-binding of essential analysis an Transcriptome is factor (2018) shock B. heat Yeast phosphorylation. Xu (1988) temperature-dependent & switchgrass H.R. in Q. Pelham network & Cai P.K. DREB2C, HSF/HSP Sorger B., of of Hu importance repressor L., the transcriptional and Lou transcripts tolerance. a Z., novel Cd VOZ1, Xie revealed (2018) root X., switchgrass Wen C.O. treated S., Lim Yuan & G., J.C. Song Arabidopsis. Hong in L.). T., responses sativa stress Kim (Oryza heat J., rice mediates Lee in are gene C., OsHsfB4b and (Hsp100) Song OsHsfA2c OsClpB (2012) cytoplasmic A. Grover Chaperones of & nutrient Stress regulation R.C. plant Cell transcriptional Mishra fine-tune M., the proteins Agarwal in D., 14-3-3 Lavania involved (2011) D., D.P. Mittal Schachtman A., & class Singh the B. of Zhang Orthologs A., rice. (2009) Basra and Y. metabolism. J.M., wheat Lee Jez & in R., E. tolerance Shin Martinoia cadmium G., confer to An WRKY1 HsfA4a Y., T., of factor Choi Lang promoter P., transcription 4031-4043. S., the Hou Lee shock G., binds J., heat Sa HSF Lee A4 N., J.U., euphratica Zhao Hwang Populus H., D., (2015) Zhang Shim S. Z., Chen Qian of M., & Ding tolerance. Introduction X. S., salt Shen (2020) Wang enhance R., A. L., Zhao Chang Jan J., F., & Sun Wang N. J., Yao Ahmed Z., tuberosum A., (Solanum Shen Iqbal potato on R.S., stress heat Khan of the effects I., plant. adverse of Munir L.) mitigates structure HsfA1d G.S., factor Solution shock Ali (1996) heat S.H., Arabidopsis’s H. Shah Ruterjans HSF24. factor & Z., transcription L. Shah heat-stress Nover tomato K.D., the Scharf P. the of U., regulates Koskull-Doring domain Gase HsfA3 von DNA-binding O., factor & Kunert transcription stress E. J., stress heat Schultheiss Vierling heat the Arabidopsis. G., and of Englich DREB2A in response factor A., genes stress transcription Ganguli of heat of E., subset cascade heat Kiehlmann A a The J., (2008) (2006) of Larkindale P. amplifier F., Koskull-Doring regulatory von Schramm a & as D. Walch Arabidopsis. stress serves in G., heat HsfA2 response Englich cytoplasmic factor E., HsfA2 in transcription Kiehlmann system: localized stress A., Hsf be Ganguli tomato may The F., and (1998) Schramm import L. Nover nuclear & efficient E. for Schmidt granules. HsfA1 R., Lyck with I., interaction Hohfeld needs H., Heider K.D., Scharf te . reik . ak .&Bul .(04 pgntcrsosst etsrs tdffrn time different at stress heat to responses Epigenetic (2014) I. RNAs. Baurle small & of involvement J. the Lamke and K., scales Brzezinka A., Stief n rncito atr eel xesv vra ewe etadnnha tesrsos pathways. response stress non-heat proteins and shock heat heat between Arabidopsis overlap of extensive profiling plants reveals Transcriptional of factors (2007) A.P. transcription response Weber and the & in M. signalling Huebner redox W.R., and of Swindell ROS subcomponent (2012) a G. Miller as stress. & response abiotic R. protein to Mittler cytosolic S., Koussevitzky The N., Arabidopsis. (2009) Suzuki in A.J. response Maule Stomatal shock of & heat Regulation F. wider (2017) Aparicio the S. R., Zhang Dreos & A., J. Sugio Xu Acid. Abscisic W., and Lukowitz Cascade Y., MPK3/MPK6 Cell Liu Pathogen-Responsive Plant T., a of Sun Functions L., Interdependent by Zhang Immunity M., Zhang J., Su o elBiol Cell Mol elSrs Chaperones Stress Cell , 29 ESLett FEBS ln elRep Cell Plant 526-542. , ln elEnviron Cell Plant , ln Sci Plant , , 17 ln o Biol Mol Plant 585 18 243-254. , 2240-2251. , , 143-147. , 37 , , 1485-1497. , 235 25 ln J Plant , 35 57-63. , 89-100. , , Cell 259-270. , 60 ln inlBehav Signal Plant , 759-772. , 53 , ln Cell Plant Planta 54 264-274. , 855-864. , 27 , 247 , 21 1439-1448. , 642-654. , , 9 e970430. , u Biochem J Eur ln Cell Plant , 236 911-921. , BMC , 21 , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. ilasB,KbaeM,BitR ika ..(00 tA7 nAaiossBcl-2-associated Arabidopsis an AtBAG7, response. protein (2010) unfolded M.B. the A in Dickman involved S is U & and Sci reticulum R. Acad endoplasmic the Britt immune in plant M., resides athanogene, regulates Kabbage 3 factor B., cassava. transcription in Williams shock signalling Heat and (2018) accumulation H. acid Shi 2209-2220. salicylic & , of C. modulation He through Y., Functional Chang response and Evolution. G., Accumulation Plant Liu Gradual with Origin, Y., Family Evolutionary Wei (2018) Gene S. Factor Xu Shock & Heat C. of Ma S., Divergence Chen Arabidopsis. X., Shi in X., tolerance Wang and heat proteins shock conferring heat bermudagrass of regulation African Transcriptional (2016) from Rep B. CtHsfA2b Huang by & J. peroxidase Liu ascorbate Z., Yang factors. W., transcription Huang stress X., heat Wang plant of diversity The (2007) Sci L. Nover Plant for Trends & required K.D. is Scharf Arabidopsis. P., H(2)O(2) Koskull-Doring in stress-induced von genes Heat shock (2006) signaling heat F. acid Schoffl of (2013) & expression abscisic K. P.M. effective Mullineaux the Shinozaki II, in & Panchuk, R.A., network S.C. Volkov Peck phosphorylation Y., protein thaliana. Ishihama a Arabidopsis MAMP-triggered J.C., in reveal pathway between Anderson phosphoproteomics Interplay F., and Takahashi (2008) Genetics F. N., Sugiyama Katagiri & T., J., Umezawa J.D. Feeney multiplexed Cohen massively A., responses. a J., Goubil defense Glazebrook CrY2H-seq: SA-mediated R., M., (2017) and Castanon Sato J.R. Ecker K., A., & Tsuda Bartlett M. mapping. Retrograde Galli J.R., interactome Z.Z., in deep-coverage Nery Zhang for Operates A., assay S.C., STN7 Huang MacWilliams (2012) R., R.M., E.M. O’Malley Chain. Garza Aro Transfer Electron & S.A., the S. Trigg in Kangasjarvi Balance Redox M., Controlling Suorsa through P.J., Signaling J. program. Jordano Gollan longevity & seed M., M. HSFA9 Ohme-Takagi Tikkanen the K., of Hiratsu function R., of Carranco Loss C., (2010) Almoguera P., Prieto-Dapena factors shock J., Tejedor-Cano heat J. by activation Jordano longevity. transcriptional & seed under synergic J.M. sunflower hinders Espinosa role in that C., involved mechanism potential Almoguera repression P., their passive Prieto-Dapena A reveal J.M., (2014) characterization Personat to Genome-wide R., (2019) (HSFs) Carranco L. J., L.). factors Tejedor-Cano Liu sativus & transcription (Raphanus L. radish shock Fan Arabidopsis Y., in heat Xie stresses of of X., abiotic site Luo analysis W., hydrolysis evolutionary Cheng GTP Y., and Wang the infection. L., in mildew Xu powdery mutation M., Tang to A response (2006) in R.W. death cell Innes enhanced 47 & confers C.A. 1E N., Frye protein Shibuya dynamin-related J., OsHsf23. S., factor, Ade transcription Toki shock D., T., heat Tang a Hagio of activation suppresses Y., via CIGR2, Fujisawa fungus protein, Biochem blast family Biotechnol B., rice GRAS with Biosci a Day inoculated for rice gene N., in elicitor-responsive death Ishii-Minami The cell (2016) N., E. Minami Hara & Y. H., Nishizawa Onodera S., Tanabe Genomics uZ,LagJ,Wn . hoX,ZogX,CoX,L . eJ iM 21)Oeepeso fTwo of Overexpression (2018) M. Yi & J. 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Hsf suite unknown a previously regulates including that L.) activator aestivum transcriptional (Triticum a wheat Bot is in Exp TaHsfA6f genes J (2015) protection C.L. stress McIntyre heat & of J. Drenth G.P., Xue nrsos oha n te ao boi tessadterrl nrglto fha hc rti genes. protein shock heat of regulation in aestivum role Triticum their from and family stresses factor abiotic shock Bot heat major The other Exp (2014) and J C.L. heat McIntyre to & response J. in Drenth S., Sadat G.P., Xue Heat Plant of Evolution into Insights New Stresses. (2020) Abiotic J. to Liu Response & in Genes D. Tea Kong of P., Analysis Expression Zhang , and X., (Hsfs) Pang Factors Shock Q., Guo P., Xu Catabolism. Proline in hn . iG,F . unS,H .&GoX 22a eoewd dnicto,transcriptome identification, Genome-wide (2020a) X. Guo & maize. D. in genes Hu temper- family S., low Hsf Duan of fruit events loquat C., lignin splicing Fu of with alternative and Regulation associated G., analysis genes (2016) Li and K.S. H., factors Chen Zhang shock & heat X.R. of Yin interaction H., involves Ge biosynthesis. lignification J., and Zhang response expression. X., ature gene HSFA7b Li regulate factor to J.K., transcription Differential motif Zeng shock E-box-like (2019) heat an Y.H. Arabidopsis to (2019) Chen binding Y. by & Wang tolerance Bot & M.T. stress Z. Geng salt stresses. Liu mediates X., abiotic Z.Q., positively Zhang and Xia J., biotic Wang C.X., under D., cassava Li Zang in P.Y., family Hou Hsf Y.H., the Hong of expression Y., Yao X.Y., system. Yu & stress-regulatory K. DREB2A Shinozaki the Y., HsfA3 Fujita of factor S., downstream transcription Commun Kidokoro cascade heat-shock J., transcriptional Arabidopsis Mizoi an the F., of in Qin analysis Functional K., (2008) Maruyama K. D., Yamaguchi-Shinozaki Todaka Y., Sakuma expression. T., gene Yoshida Arabidopsis shock-responsive heat (2011) in K. regulators Yamaguchi-Shinozaki Genomics positive M., & Genet main Seki K. the J.M., Mol Shinozaki as Kim F., function K., factors Schoffl Maruyama transcription K., Y., Nakashima HsfA1 Sakuma J., Y., Mizoi Osakabe S., Kidokoro D., transgenic Todaka J., Expression in Nakajima (2008) N., stresses K. Ohama environmental T., Oda to Yoshida & tolerance M. enhances Iwabuchi OsHsfA2e H., factor Hirochika transcription Arabidopsis. M., stress Matsui heat Y., rice Shock of Kondou Heat T., of Ichikawa Investigation N., Genome-Wide Development. Yokotani (2020) Anther X. in Roles Song Possible & Sci and L. L.) Mol Zhang aestivum W., J evolution (Triticum Li Adaptive Wheat (2014) Q., in C. Family Liu Factor Xu G., Transcription grasses. & Hu G. in X., Liang Yang factors Y., J., transcription Yuan Ye Y., stress encodes Hu heat Spl7, E., of gene, Zhang expression Y., leaf divergent Zhou spotted and Y., rice Gao A Y., Wang (2002) Z., K. Yang protein. Yamada the factor & for transcription M. receptor stress Ashikari second heat Arabidopsis. H., a a in Lin is responses M., PEPR2 Yano defense (2010) U., to C.A. Yamanouchi contributes Ryan and & peptides F.E. Pep2 Tax and A.C., Pep1 Bryan A., Huffaker Y., stresses Yamaguchi photooxidative to response in involved factors Arabidopsis. transcription shock in heat of Functions (2016) Y. Yabuta 9 . , 70 5355-5374. , , 368 , , , 21 66 65 Planta ln elEnviron Cell Plant 515-521. , . 1025-1039. , 539-557. , isiBoeho Biochem Biotechnol Biosci , , 227 286 x Bot Exp J 957-967. , 321-332. , , , 39 69 1780-1789. , 2005–2021. , rcNt cdSiUSA S U Sci Acad Natl Proc , 80 1254-1263. , 29 , c Rep Sci 99 M vlBiol Evol BMC Genome 7530-7535. , , ln Cell Plant 10 , 8073. , 62 ice ipy Res Biophys Biochem 563-569. , , , 14 22 147. , lns(Basel) Plants 508-522. , Exp J Int Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis protein thaliana HSF Arabidopsis thaliana Arabidopsis Species Le- O’Donnell, Provart, 2019). Kolas, Tyers, & Chang, & Usadel Dolinski Boucher, Chatr-Aryamontri, Grene, Rust, Coulombe-Huntington, Brady, Breitkreutz, Willems, Dolma, Tyers, Stark, Zhang, Oughtred, Dolinski, McAdam, 2016, Oughtred, ung, Orchard, Pasha, & Gaudinier, Licata 2019, Nahal, Sivieng, Wu, ban, Gene OsHSFA3 Rice (2020) Y.M. Lv & ROS B. and Zhou Acid S.J., Abscisic Factor Tang on Shock Depending K., Biosynthesis Heat Zhou Polyamine (2018) Modulating Levels. D.J., J.Q. by Yu Gao Tolerance & Drought M., Production(1). Improves Y.H. H2O2 Zhang Zhou Triggers M.D., K., and Zhu Shi Resistance Facilitates X.J., SlSIZ1 Nematode Ligase Xia Gene-Mediated E3 L.L., R SUMO Yin Physiol for (2018) J.J., Essential Q. Cao Is Meng X.C., HsfA1a & Xu N. Ma J., Y., Zhou Wang subcellular Arabidopsis. Z., in and Tomato. Liu stress in dynamics J., Tolerance heat Lv mitochondrial Heat by S., of Wang caused Characterization S., shock damage Zhang (2009) oxidative heat C. alleviates the Gao HsfA2 of that & Bot profiling reveal Exp D. expression ROS Xing and of Y., analysis localization L.). 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Brassica Zhou from Y., factor Liu scription Y., Wang X., Zhu 471-480. , n o Sci Mol J Int , , 176 60 oplto fHFitrcin ndffrn ln pce.Poenpoenitrcin of interactions Protein-protein species. plant different in interactions HSF of Compilation 2073-2091. , 2456-2471. , SAAGF 433poenA-S(Shin (Shin al., et (Shin, AC-MS binding HSF AC-MS protein 14-3-3 AC-MS protein 14-3-3 HSBP protein 14-3-3 GRF8 down, FRET, pull Y2H, HSFA1A GRF3 kinase kinase protein protein HSFA1A GRF1 CBK3 HSFA1A CRK1, HSFA1A CDC2a HSFA1A HSFA1A , 21 . ln elPhysiol Cell Plant rti aeoyTcnlg Reference Technology Category protein Partner hlaa)(rbdpi,21,Dn,Lu og eul,Este- Ierullo, Song, Lau, Dong, 2011, (Arabidopsis, thaliana/) , 59 58-71. , 30 Nature protein , 428 764-767. , PeerJ , 8 e8467. , PH(s,Li& Lai (Hsu, PP2H phosphorylation phosphorylation LSONE PLoS Cell in 2010) Jinn, 2011) 2011) 2011) 2008) al., et (Liu, 1997) sl., et (Reindl, , Cell Plant 125 , 13 tal. et al. et J , , , , , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis protein HSF thaliana Arabidopsis Species SABHF1 F S iC(Yoshida BiFC BiFC, AC-W, HSF TF, (Czarnecka- HSF HSFA1D TF, down pull binding HSF HSFA1B HSFA1B TF general down, pull (Liu HSBP HSFA1B TF general TFIIB HSFA1B Y2H (Yoshida kinase protein HSFA1A TBP2 protein (Yoshida TBP1, & (Kim pathogen & HSFA1A TAGK2 (Kim BiFC sphingosine HSFA1A PP7 BiFC sphingosine HSFA1A NPR1 Y2H al., et Y2H (Li, HSP HSFA1A LCBK2 (Yoshida HSP HSFA1A LCBK1 Y2H BiFC, HSP HSFA1A HSP90-3 HSP BiFC HSFA1A HSP90-1 HSF TF, HSFA1A HSP70-1 BiFC, AC-W, HSF HSFA1A HSP70 TF, (Hsu HSFA1A HSFA2 Y2H HSF BiFC, HSFA1A HSFA1D TF, HSFA1A HSF HSFA1A HSFA1B TF, HSFA1A HSFA1A HSFA1A TFIID1 Reference Technology Category protein Partner 31 protein down, pull TF general phosphatase defense kinase kinase uldw,P2 Dn,e al., et (Dong, PP2H down, pull al., et (Liu, EMSA Y2H, (Liu al., et (Olate, Y2H (Liu Co-IP BiFC, Y2H Y2H Y2H EMSA Y2H 2010) 09 Hsu 2019, 1998) Schoffl, & (Reindl 2007) 2018) 2008) 2008) al. al. al. al. 2008) 2011) oshida Y 2011) Yoshida Li 2004) Verner 2002) Schoffl, 2002) Schoffl, 2010a) Li 2011) al., et Yoshida, 2010b, al., et Li, (Hsu 1998) Schoffl, & Reindl 2004, al., et Verner, (Czarnecka- (Hsu 2011) , 2011) , 2011) , 2011) , tal. et tal. et tal. et al. et tal. et tal. et tal. et al. et tal. et 2010b, , 2010b, , tal. et et et et et tal. et , , , 2010, , 2010, , 2010, , tal. et , , , , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis protein thaliana HSF Arabidopsis thaliana Arabidopsis Species SAEbI6T,bI r2-e (Trigg CrY2H-seq (Trigg (Trigg bZIP CrY2H-seq TF, CrY2H-seq Homeobox TF, BR hormone, (Trigg bZIP6 chromatin BLH7 (Trigg CrY2H-seq (Yoshida TF, BES1 HSFA1E (Yoshida CrY2H-seq ATXR5 HSFA1E ZnF pathogen TF, AGL39 HSFA1E BiFC bHLH TF, (Ohama HSFA1E AT5G05120 BiFC DNA (Ohama chromatin, HSFA1E NPR1 LC-MS/MS (Ohama HSP HSFA1D MUTE LC-MS/MS (Trigg HSP HSFA1D MBD10 LC-MS/MS HSP HSFA1D HSP90-3 CrY2H-seq HSP HSFA1D HSP90-1 HSP HSFA1D HSP70-4 BiFC, HSF TF, (Trigg HSFA1D HSP70-3 (Trigg HSFA1D (Yoshida HSP70-1 HSF CrY2H-seq TF, HSFA1D (Yoshida HSFA9 CrY2H-seq VP1-B3 Homeobox TF, HSFA1D TF, HSFA1D BiFC HSFA1D al., et (Trigg, AT3G19070 signaling, BiFC pathogen (Li HSFA1D ARF2 CrY2H-seq HSP HSFA1D ANNAT4 Y2H BiFC, HSP HSFA1D NPR1 HSF TF, HSFA1D HSP90-3 HSF TF, HSP90-1 HSFA1B HSFA1B HSFA9 HSFA1B HSFA2 HSFA1B HSFA1B rti aeoyTcnlg Reference Technology Category protein Partner 32 structure MADS-box defense methylation Ca2+ defense r2-e (Trigg (Trigg CrY2H-seq CrY2H-seq (Olate Co-IP BiFC, (Trigg CrY2H-seq et (Ohama, (Olate LC-MS/MS Co-IP BiFC, LC-MS/MS 2017) 2017) 2018) 2017) 2018) l,2016) al., al. al. al. al. 2010a) 2017) 2017) 2017) 2017) 2017) 2016) 2016) 2016) 2017) 2017) 2017) 2017) (Ohama al. Yoshida 2016, 2011) , 2011) , 2011) , 2011) , 2011) , tal. et tal. et al. et al. et tal. et al. et al. et al. et al. et al. et al. et al. et tal. et al. et , tal. et al. et al. et tal. et et et et et , , , , , , , , , , , , , , , , , et Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis protein thaliana HSF Arabidopsis thaliana Arabidopsis Species SA SC F S r2-e (Trigg (Trigg CrY2H-seq (Trigg (Trigg CrY2H-seq HSF TF, CrY2H-seq CrY2H-seq HSF TF, (Trigg HSF (Trigg HSFC1 TF, ZnF TF, CrY2H-seq HSFA7A chromatin CrY2H-seq HSFA6A HSFA3 bHLH TF, GAL2 bHLH HSFA3 TF, & BRM (Meiri al., splicing et HSFA3 RNA, (Evrard, BNQ3 2017) (Li, HSFA3 chromatin BIM1 down HSFA3 pull BiFC protein ATO HSFA3 kinase protein ADA2A Y2H HSFA3 BiFC, SUMO1 Y2H, HSFA3 HSP MPK6 HSF TF, HSFA3 HSF (Dong TF, HSFA2 HSP90-1 binding HSF HSFA2 HSFA3 prediction HSFA2 HSFA2 HSBP HSFA2 hormone (Trigg HSFA2 FKBP62, HSFA2 CrY2H-seq SMAD/FHA protein AHK3 (Trigg pathogen AT3G07260 HSFA2 RAV CrY2H-seq TF, PIAL2 HSFA2 NPR1 HSFA1E DOF repeat TF, WD40 NGA1 HSFA1E MUM1 LUH, HSFA1E COG1 HSFA1E HSFA1E HSFA1E ROF1 Reference Technology Category protein Partner 33 remodeling factor remodeling sumoylation protein protein repeat tide tetratricopep- memory, HS protein sumoylation defense protein 2 Ern,e al., et (Efroni, Y2H (Trigg (Trigg CrY2H-seq (Cohen-Peer, CrY2H-seq BiFC Y2H, (Hsu & (Meiri PP2H BiFC (Trigg (Trigg CrY2H-seq (Olate CrY2H-seq Co-IP BiFC, (Trigg CrY2H-seq cross-linking 2017) 2017) 2017) 2017) 2018) 2017) 2013) 2010) al., et 2010) 2009) Breiman, 2017) 2017) 2017) 2017) 2017) 2017) 2013) 2009) Breiman, 2019) 2017) 2017) aua,2011) Sakurai, & (Enoki tal. et tal. et al. et al. et al. et al. et tal. et al. et al. et al. et al. et al. et al. et al. et tal. et tal. et , , , , , , , , , , , , , , , , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis protein thaliana HSF Arabidopsis thaliana Arabidopsis Species SACbI5 F ZPCYHsq(Trigg (Trigg (Trigg CrY2H-seq CrY2H-seq (Trigg ERF/AP2 TF, CrY2H-seq (Trigg bZIP TF, (Dong CrY2H-seq DEWAX bZIP CrY2H-seq TF, prediction bZIP52 bHLH TF, VP1-B3 TF, HSFA4C At1g35490 transport Y2H, AC-W, HSFA4C BHLH118 (Baniwal protein AC-W, kinase HSFA4C ARF11 protein kinase et HSFA4C VPS55 protein (Baniwal, Y2H, AC-W, HSFA4C Y2H kinase MPK6 protein HSFA4A (Perez-Salamo, MPK4 Y2H HSF TF, HSFA4A Y2H FRET, MPK3 HSFA4A HSF TF, HSFA5 (Trigg HSF HSFA4A TF, TF, HSFA4C CrY2H-seq chromatin HSFA4A HSFA4A chromatin HSFA4A AGL29 S1FA TF, SWI3C (TriggHSFA4A transcription, SWI3B HSFA4A transcription, (Trigg CrY2H-seq S1FA2 HSFA3 translation (Trigg OFP18 HSFA3 CrY2H-seq NAC TF, OFP13 HSFA3 CrY2H-seq methyltransferase NOT9B HSFA3 IAA TF, NAC066 HSFA3 LSMT-L HSFA3 IAA33 HSFA3 HSFA3 HSFA3 rti aeoyTcnlg Reference Technology Category protein Partner 34 MADS-box remodeling remodeling OVATE OVATE regulation rdcin(Dong (Efroni (Efroni prediction Y2H Y2H (Trigg (Trigg CrY2H-seq (Trigg CrY2H-seq CrY2H-seq phosphorylation FRET, kinase phosphorylation FRET, 2019) 2017) 2017) 2017) 2013) 2013) study al. 2017) 2017) 2017) 2017) 2017) 2019) study this 2007), al., 2014) al., et 2017) 2017) 2017) 2017) tal. et (Perez-Salamo 2019) al., et (Andrasi, al. et (Perez-Salamo 07,this 2007), , 2014) , 2014) , tal. et al. et al. et tal. et al. et al. et al. et al. et al. et al. et al. et al. et tal. et tal. et tal. et al. et et , , , , , , , , , , , , , , , , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana protein Arabidopsis HSF thaliana Arabidopsis Species SA I1T,bL r2-e (Trigg CrY2H-seq (Shin bHLH TF, (Shin ZnF TF, AC-MS binding HSF CIB1 (Trigg AC-MS protein 14-3-3 (Trigg BBX9 protein 14-3-3 CrY2H-seq HSBP CrY2H-seq (Trigg HSFA9 al., et (Hwang, GRF8 Homeobox TF, HSFA9 TCP TF, CrY2H-seq (Trigg GRF3 HSFA8 (Trigg translation HSFA8 WOX13 CrY2H-seq Y2H HSF TF, HSFA8 TCP2 CrY2H-seq HSFA7A NOT9B bZIP TF, ZnF TF, (Trigg HSFA7A HSFC1 TCP TF, HSFA7A At1g35490 CrY2H-seq chromatin (TriggHSFA7A VOZ1 chromatin (Trigg HSFA7A TCP4 HSF CrY2H-seq TF, HSFA6A SWI3C chromatin CrY2H-seq (Dong HSFA6A SWI3B et (Popescu, NAC TF, HSFA6A HSFC1 phosphorylation MYB prediction TF, (Trigg HSFA6A BRM kinase (Trigg protein NAC062 HSFA6A (Baniwal CrY2H-seq transport (Trigg MYB56 CrY2H-seq HSFA6A MPK4 HSFA5 CrY2H-seq NAC TF, MYB MATE TF, HSFA5 Y2H MYB TF, HSFA5 At5g18037 HSFA5 MYB75 HSF TF, HSFA4C MYB117 HSFA4C HSFA5 HSFA4C HSFA4C rti aeoyTcnlg Reference Technology Category protein Partner 35 B-Box protein control remodeling remodeling remodeling r2-e (Trigg (Dong CrY2H-seq prediction (Trigg CrY2H-seq (Efroni (Efroni Y2H (Efroni Y2H Y2H tal. et al. 2017) 2019) 2017) 2013) 2013) 2013) 2017) 2011) 2011) 2017) 2017) 2017) 2017) 2014) 2017) 2017) 2017) 2017) 2009) al., 2019) 2017) 2017) 2017) 07 Trigg 2007, , 2017) , tal. et al. et tal. et al. et tal. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et tal. et tal. et tal. et al. et al. et et , , , , , , , , , , , , , , , , , , , , , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis protein thaliana HSF Arabidopsis thaliana Arabidopsis Species SBBLC5pooytei 2 (Li (Li Y2H Y2H photosynthesis HSF TF, enzyme, LHCA5 chromatin HSFB2B repeat WD40 HEMC HSFB2B repeat WD40 FGT1 HSFB2B repeat WD40 TPR4 HSFB2B repeat WD40 HSFB2B TPR3 repeat WD40 HSFB2A TPR2 repeat WD40 (Li TPR4 HSFB2A repeat WD40 TPR3 (Trigg repeat WD40 HSFB2A (Trigg TPR2 HSFB1 Y2H, (Li CrY2H-seq enzyme, TPR1 CrY2H-seq HSFB1 ERF/AP2 (Trigg TF, TPL HSFB1 HSF TF, PYRP2 ZnF HSFB1 CrY2H-seq TF, Y2H ORA47 HSFB1 (Trigg HSFB1 bHLH TF, HSFB1 chaperon FES1 CrY2H-seq HSFB1 death, (Popescu cell BHLH010 HSFB1 phosphorylation (Trigg (Trigg NFYC TF, ATJ3 (Trigg HSFB1 kinase protein APG8H CrY2H-seq CrY2H-seq HSFB1 CrY2H-seq NF-YC1 Homeobox HSFB1 TF, ERF/AP2 TF, MPK6 EIN3 TF, HSFB1 At2g02060 HSFA9 ERF112 HSFA9 EIL2 HSFA9 HSFA9 HSFA9 rti aeoyTcnlg Reference Technology Category protein Partner 36 plastid remodeling protein protein protein protein protein protein protein protein plastid autophagy 2 (Li (Causier (Trigg Y2H (Causier CrY2H-seq (Causier Y2H (Causier Y2H (Causier Y2H (Causier Y2H (Causier Y2H et (Causier, Y2H (Li Y2H Y2H Y2H (Li Y2H 2010b) 2017) 2012) 2012) 2012) 2012) 2012) 2012) 2012) 2010b) 2010b) l,2012) al., 2010b) 2010b) 2010b) 2010b) al. 2017) 2017) 2017) 2017) 2017) 2017) 2017) 2009) , tal. et al. et al. et tal. et al. et al. et al. et tal. et tal. et al. et al. et al. et al. et al. et al. et , , , , , , , tal. et al. et al. et al. et al. et al. et al. et et , , , , , , , , , , , , , , , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis protein thaliana HSF Arabidopsis thaliana Arabidopsis Species SB C1 F C r2-e (Trigg (Trigg CrY2H-seq CrY2H-seq (Trigg TCP TF, CrY2H-seq TCP TF, (Trigg (Trigg TCP14 HSF CrY2H-seq TF, RING/U-box CrY2H-seq TCP10 ERF/AP2 TF, AT3G43430 TCP HSFB3 TF, (Trigg REN1 HSFB3 translation RAP2.5 CrY2H-seq HSFB3 (Trigg PTF1 Homeobox ZnF- TF, HSFB3 TF, (Trigg NOT9B ZnF- HSFB3 TF, CrY2H-seq (Trigg At4g03250 CrY2H-seq (Trigg HSFB3 Homeobox TF, HB30 CrY2H-seq HSFB3 trihelix (Trigg TF, CrY2H-seq HB21 (Trigg HSFB3 GT3A binding RNA CCT HSFB3 TF, CrY2H-seq CrY2H-seq ENAP1 HSFB3 EMB1967 bHLH TF, HSFB3 trihelix TF, CIA2 HSFB3 TF, BHLH010 HSFB3 TF, ASIL2 HSFB3 repeat WD40 AGL17 HSFB3 (Li repeat WD40 AGL16 HSFB3 repeat WD40 TPR4 HSFB3 repeat WD40 Y2H TPR3 HSFB3 photosynthesis TPR2 HSFB2B TPL HSFB2B PSBP-1 HSFB2B MUSE14, HSFB2B HSFB2B HSFB2B rti aeoyTcnlg Reference Technology Category TRAF1A protein Partner 37 protein control Homeobox Homeobox MADS-box MADS-box protein protein protein protein protein domain TRAF r2-e (Trigg CrY2H-seq (Trigg CrY2H-seq (Trigg (Trigg CrY2H-seq CrY2H-seq (Trigg (Causier (Trigg CrY2H-seq (Causier CrY2H-seq (Causier Y2H (Causier Y2H Y2H (Li Y2H Y2H 2017) 2017) 2017) 2017) 2017) 2017) 2012) 2012) 2012) 2012) 2010b) 2010b) 2017) 2017) 2017) 2017) 2017) 2017) 2017) 2017) 2017) 2017) 2017) 2017) tal. et tal. et tal. et al. et al. et al. et al. et al. et tal. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et , , tal. et al. et al. et al. et , , , , , , , , , , , , , , , , , , , , , , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. lcn a SB FI eea Fpl on(Czarnecka- down pull TF General TFIIB annuus Helianthus HSFB1 max Glycine japonica Eriobotrya thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis thaliana Arabidopsis protein thaliana HSF Arabidopsis thaliana Arabidopsis Species SAAHF9T,HFYH iC(Tejedor-Cano, BiFC Y2H, al., et (Zeng, HSF TF, (Trigg (Trigg BiFC Y2H, (Trigg ERF/AP2 TF, CrY2H-seq CrY2H-seq HSFA9 (Trigg CrY2H-seq (Trigg ZnF bZIP TF, CrY2H-seq TF, AP2-1 (Trigg HSFA4A TCP TF, CrY2H-seq AT2G47850 CrY2H-seq TCP (Arabidopsis, TF, TGA9, (Trigg HSF3 TCP TF, TCP4 TCP HSFC1 TF, (Trigg CrY2H-seq TCP14 HSFC1 Y2H TCP10 CrY2H-seq PLATZ TF, HSFC1 kinase protein PTF1 HSFC1 NFYB (Trigg TF, At2g27930 HSFC1 (Arabidopsis, PBL35 HSFC1 CrY2H-seq chromatin (Trigg NF-YB7 ZnF- HSFC1 TF, bHLH TF, HSFC1 CrY2H-seq Y2H (Trigg MBD6 HSFC1 HB30 cytomatrix TCP CrY2H-seq TF, FIT1 HSFC1 ABA hormone, (Trigg At1g19980 (Trigg HSFC1 death, cell (Trigg BRC1 HSFC1 CrY2H-seq CrY2H-seq chromatin ARIA HSFC1 CrY2H-seq Homeobox TF, AMC1 HSFC1 TCP TF, ADA2A HSFC1 TCP TF, WOX13 HSFC1 TCP9 HSFC1 TCP4 HSFB3 HSFB3 HSFB3 rti aeoyTcnlg Reference Technology Category protein Partner 38 methylation DNA structure, Homeobox metacaspase remodeling r2-e (Trigg (Trigg CrY2H-seq CrY2H-seq (Trigg (Trigg CrY2H-seq CrY2H-seq 2017) 2017) 2017) 2017) ta. 2014) al., et 2004) Verner 2016) 2017) 2017) 2017) 2017) 2017) 2017) 2017) 2011) 2017) 2017) 2011) 2017) 2017) 2017) 2017) 2017) tal. et al. et al. et al. et tal. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et tal. et , , , , , , , , , , , , , , , , , , , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. rz aiaHf4 SBcT,HFYH(Mittal lycopersicum (Mittal Solanum Y2H lycopersicum (Mittal Solanum cross-link, (Mittal lycopersicum HSF Solanum TF, Y2H lycopersicum Solanum HSF TF, (Singh lycopersicum Y2H Solanum Y2H HSFB4c HSF TF, lycopersicum BiFC Y2H, al., Solanum et HSFB4b (Singh, lycopersicum HSF TF, Solanum HSF cross-link, TF, lycopersicum HsfB4b HSFA7 BiFC Y2H, Solanum HsfB4b HSFA2a HSP sativa Oryza HSF TF, cross-link, HSF26 sativa Oryza HsfB4b ClpB-cyt HSP HsfB4b sativa Oryza HSF HSFA9 TF, HsfB4b sativa Oryza ClpB-cyt HsfB4b sativa Oryza HSFA2c HSFA9 sativa Oryza HSFA7 sativa Oryza HSFA2c sativa Oryza sativa Oryza peruvianum Lycopersicon peruvianum Lycopersicon peruvianum Lycopersicon protein longiflorum HSF Lilium annuus Helianthus Species SA S9 S 2 (Hahn (Hahn BiFC, Y2H Y2H, (Hahn Y2H al., et (Hahn, HSF TF, HSP Y2H HSP complex Y2H, Y2H al., et HSFA5 (Zhang, HSP90 small HSP, HSP BiFC Y2H, HSP70 HSFA4b HSP Hsp17.4-CII HSFA2 Sumoylation HSP90 HSFA2 HSP70 HSFA2 chromatin SlSIZ1 HSFA1a HAC1/CBP HSFA1a HsfA1 HsfA1 pull-down, (Chan- HSF chromatin al., TF, et (Gong, Co-IP Y2H, HAC1/CBP BiFC Y2H, HSFB1 HSF TF, HSFB1 HSF TF, HSFA1 Aux/IAA HSFA2 HSFA2 HSFA1 IAA27 HSFA1 HSFA4A rti aeoyTcnlg Reference Technology Category protein Partner 39 structure structure protein EMSA pull-down, EMSA pull-down, (Tejedor-Cano BiFC pull-down form BiFC BiFC BiFC EMSA 2004) 2004) 2011) 2011) 2011) (Bharti (Bharti al. et 2011) al. Singh 2011, 2011) 2011) 2011) 2018) 2012) 2012) 1998) al., et Scharf, 2004, al., et Port, 2009, al., et Schaminet, 2014) al. (Baniwal 2004) (Port 2011) (Mittal 2011) (Mittal 2011) al., et (Mittal, 2004) al., et (Bharti, 2012) , 2007) , 2014) , tal. et tal. et al. et al. et tal. et tal. et al. et al. et al. et tal. et al. et tal. et al. et et , , , , et , , , , , , , , , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. O4A3105ncesAlne itn iepoen1. 445513.1 5.6 19.6 13.2 3.8 5.4 70.2 5.5 3.5 10.3 5.7 2.8 10.9 2.8 14.4 3.4 94.8 6.8 3.0 23.4 14.1 % Coverage 4.5 6.4 68.2 Count 10.3 Peptide 4.7 5.2 % Coverage 3.8 13.3 3.7 Count Peptide 6.5 13.6 103.0 3.7 3.2 2.8 7.1 protein like histone linker A 2 protein organizing Hsp70-Hsp90 6.5 7 4.3 1 regulator coverage 4.0 chaperone chloroplast nucleus molecular Reduced 3 family resistance FIGURES BAG disease AT3G18035 nucleus Enhanced nucleus AT1G62740 ER, 8 factor nucleus AT5G62390 protein Pre-mRNA-processing-splicing transduction signal HON4 AT1G01320 cytosol TIM-barrel Name Protein HOP2 AT3G60190 HSFA4C HSFA5 BAG7 nucleus REC1 location cytosol AT1G80070 HSFA4A AT5G66420 EDR3 nucleus nucleus TIM-barrel AT5G45710 AT4G13980 PRP8 nucleus AGI HSFA4C AT4G18880 HSFA5 HSFA4A Gene binding spectrometry. HSF mass which %). by experiment, Coverage identified each count, were in (Peptide HSFA4A-YFP used samples binding with were 6 EMP2 HSF lines co-purified of HSBP1, independent binding averages Proteins 150 Two show HSF without Numbers times. extraction. or protein three with prior treated repeated were hours were promoter 6 pHSFA4A for of EMP2 control NaCl HSBP1, the mM under fusion HSFA4A-YFP the bindingpressing HSF 2. HSBP2 HSFA5 Table binding EMP2 HSF HSBP1, binding HSF HSFA4d EMP2 HSBP1, mays HSFA4a Zea HSBP2 mays HSFA3 Zea mays HSFA2e Zea mays HSFA2c Zea mays Zea mays Zea thaliana Arabidopsis salsuginea, Thellungiella thaliana Arabidopsis salsuginea, Thellungiella protein lycopersicum HSF Solanum lycopersicum Solanum Species dnicto fHF4 neatn rtis w ek-l rngncAaiosspat,ex- plants, Arabidopsis transgenic weeks-old Two proteins. interacting HSFA4A of Identification sAdHP03HPBF (Higashi et (Higashi, BiFC (Hahn (Hahn BiFC HSP Y2H Y2H HSP HSP90-3 HSP HSP HSP90-1 HsfA1d HSP90 HSP70 HsfA1d HsfB1 HsfB1 rti aeoyTcnlg Reference Technology Category protein Partner 40 protein protein protein protein protein protein 2 (Fu (Fu (Fu Y2H (Fu Y2H (Fu al., Y2H et (Fu, Y2H Y2H Y2H oto oto attetetSl treatment Salt treatment Salt Control Control 2013) 2017) al., et Roth, 2011, 2011) 2006) 2006) 2006) 2006) 2006) 2013) al., 2006) tal. et al. et al. et al. et al. et tal. et al. et tal. et , , , , , , , , Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. dnicto fHF4-neatn rtisb omupeiiainadms spectrometry. mass and coimmunprecipitation by proteins HSFA4A-interacting of Identification Methods Supplemental S4: Table S3: Table S2. plants. Table overexpressing transgenic or mutants HSF of negative and S1. Table positive indicate arrows or data lines Supplemental red and Green WRKY30 2004). or respectively. ZAT12 Mittler, regulation, as & such TFs Liang other Davletova, or (Rizhsky, proteins (Baniwal protective HSFA4A encode which repress genes, (Davletova target can of which number HSFA5, a activate and HSFA4C with (Andrasi heteromers the multimerisation to bind promotes can al. which Galli, TFs HSFA4A, Huang, WRKY phosphorylate O’Malley, and PEPR1, (Bartlett, MPK6 MYB kinases study HB, during receptor DAP-seq ZAT, the generated HSF, a by (Andrasi 2017), to is mediated Ecker, According ROS are & Gallavotti EDS4. signals Nery, FLG22 PCD-regulating pathogens. of and the various Pep2 Expression and EF-Tu, by PEPR2, enhance microbial networks. and can that regulatory suggest signals conditions data other environmental stress with biotic extreme which, and of stress, number abiotic up in a indicates and HSFA4A green by agents and oxidative Red 5. or Genevestigator. elicitors Figure from with compiled treated were downregulation, pathogens, respectively. Data or with downregulation, mutants. up or infected redox-related indicate plants or boxes in defense green expression in from and HSF Red compiled and 4. boxes. genes, ecotypes Figure color type HSF the wild Arabidopsis of including of side genotypes right expression while the the respectively. side in left on green listed the treatments and are in mutants red stress indicated microarray- codes abiotic are of Treatments color of hundreds In Genevestigator. Effect of germination. average 3. during is Figure levels value respectively. transcript Each downregulation, in or Change levels. up B) indicates transcript higher data. developmental expression indicates different derived color in Darker levels stress Transcript in heat A) stages. genes after (https://genevestigator.com). HSF or database of plates Genevestigator control from regulation salt Developmental on after similar rates 2. was higher Figure plants at of survived survival and shown). and affected (Andrasi (not Growth less recovery only were stresses. for plants days combined overexpressing (37 10 the HSFA4A temperature for and that high medium Note, with growth plantlets germinated treated 2019). standard vitro and to , in NaCl transferred days-old mM 10 subsequently 100 were stress. containing Plantlets heat (HSFox1, and medium plants salt Arabidopsis to overexpressing of transferred HSFA4A combination and were or (Col-0) salt type to wild exposed of lines) HSFox2 survival and Growth 1. Figure 04.RS npriua H particular in ROS, 2014). , tal. et ucinldvriyo ln S ee.Ls fpbiain eotn tesrltdphenotypes stress-related reporting publications of List genes. HSF plant of diversity Functional rncitpolso rbdpi etsokgnsi epnet ahgn n elicitors. and pathogens to response in conditions. genes stress shock abiotic heat to Arabidopsis response of in profiles genes Transcript thaliana. shock Arabidopsis heat of Arabidopsis ecotypes of 34 profiles in Transcript genes HSF 21 of levels Transcript tal. et 09.HF4 a neatwt aiu rtis(e:Tbe1 ) P3 P4and MPK4 MPK3, 2). 1, Table (see: proteins various with interact can HSFA4A 2019). , 05 Perez-Salamo 2005, , 2 O 2 a tblz rmr (Perez-Salamo trimers stabilize can tal. et HSFA4A 04.TeeTscnidc nte e ftre genes target of set another induce can TFs These 2014). , 41 rbdpi thaliana Arabidopsis xrsin(Perez-Salamo expression tal. et rncitdt eecompiled were data Transcript . tal. et tal. et 04.HF4 a form can HSFA4A 2014). , tal. et 09 Perez-Salamo 2019, , 07.HF4 can HSFA4A 2007). , HSFA4A 04.Transcript 2014). , HSFA4A o )fr4days. 4 for C) sinduced is promoter tal. et et Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. 42 Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. 43 Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. 44 Posted on Authorea 20 Jul 2020 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.159527034.41187970 — This a preprint and has not been peer reviewed. Data may be preliminary. 45