The bHLH transcription factor CsPIF4 positively regulates high temperature-induced hypocotyl elongation in cucumber

High temperature-induced hypocotyl elongation is a typical thermomorphogenesis trait that may significantly affect early seedling growth and subsequent crop yield.The ambient temperature and endogenous auxin are two critical factors that regulate hypocotyl growth.However,the mechanism of temperature and auxin integration in horticultural plants remains poorly understood.In this study,the roles of the basic helix-loop-helix transcription factor CsPIF4 in regulating auxin biosynthesis genes and the auxin content in the hypocotyl of cucumber(Cucumis sativus L.)seedlings under high temperature were investigated.qRT-PCR and in situ hybridization analysis revealed that expression of CsPIF4 was enhanced in the epidermis and vascular bundles in the hypocotyl of cucumber seedlings in response to high temperature.qRT-PCR and HPLC analysis showed that CsPIF4 positively regulated transcription of the auxin biosynthesis gene CsYUC8 and the auxin content in the hypocotyl under high temperature(35℃).The CRISPR/Cas9-mediated knockout of CsPIF4 resulted in a shorter hypocotyl compared with that of the wild type,together with decreased expression of CsYUC8 and lower auxin content in response to high temperature.Furthermore,biochemical assays showed that CsPIF4 could bind directly to the G-box motif of the CsYUC8 promoter and thereby activate CsYUC8 expression.These findings provide insight into the molecular mechanism of high temperature-mediated hypocotyl elongation in cucumber.

Influence of K + on the Coupling Between ATP Hydrolysis and Proton Transport by the Plasma Membrane H +_ATPase from Soybean Hypocotyls

The plasma membrane vesicles were purified from soybean (Glycine max L.) hypocotyls by two_phase partitioning methods. The stimulatory effects of K + on the coupling between ATP hydrolysis and proton transport by the plasma membrane H +_ATPase were studied. The results showed that the proton transport activity was increased by 850% in the presence of 100 mmol/L KCl, while ATP hydrolytic activity was only increased by 28.2%. Kinetic studies showed that K m of ATP hydrolysis decreased from 1.14 to 0.7 mmol/L, while V max of ATP hydrolysis increased from 285.7 to 344.8 nmol Pi·mg -1 protein·min -1 in the presence of KCl. Experiments showed that the optimum pH was 6.5 and 6.0 in the presence and absence of KCl, respectively. Further studies revealed that K + could promote the inhibitory effects of hydroxylamines and vanadates on the ATP hydrolytic activity. The above results suggested that K + could regulate the coupling between ATP hydrolysis and proton transport of the plasma membrane H +_ATPase through modulating the structure and function of the kinase and phosphatase domains of the plasma membrane H +_ATPase.

Effect of Lysophosphatidylcholine on ATP and ρ-Nitrophenyl Phosphate Hydrolysis by the Plasma Membrane H^+-ATPase from Soybean Hypocotyls

The stimulatory effect of lysophosphatidylcholine (lyso_PC) on ATP and ρ_nitrophenyl phosphate (PNPP) hydrolysis by the plasma membrane H +_ATPase from soybean (Glycine max (L.) Merr.) hypocotyls was studied. Results showed that lyso_PC stimulated the hydrolysis of ATP; ATP hydrolysis was enhanced dramatically when lyso_PC was within 0-0.03%, and increased slightly when lyso_PC was higher than 0.03%. At the concentration of 0.03%, lyso_PC stimulated ATP hydrolysis by 80.5%. Kinetics analysis showed that V max increased from 0.46 μmol P i·mg -1 protein·min -1 to 0.87 μmol P i·mg -1 protein·min -1 while K m increased from 0.88 mmol/L to 1.15 mmol/L under lyso_PC treatment. The optimum pH of ATP hydrolysis was shifted from 6.5 to 7.0 . Moreover, it was found lyso_PC enhanced the inhibition of ATP hydrolysis by hydroxylamine. In the presence of 200 mmol/L hydroxylamine, ATP hydrolysis was inhibited by 74.4%, while it was inhibited by 84.4% when treated with lyso_PC. However, PNPP hydrolysis and the inhibitory effect of vanadate were not affected by lyso_PC. The above results indicated that the kinase domain might be an action site or regulatory region of the C_terminal autoinhibitory domain in the plant plasma membrane H +_ATPase.

Effects of Natural Brassinolide on Seed Germination and Hypocotyl Elongation of Beans

Seeds of soybeans and mung beans were soaked into five different concentrations （0, 0.05, 0.10, 0.20, 0.30 mg/L） of natural brassinolide （ NBR ） solution. According to the results, natural brassinolide treatment could improve seed germination rate and hypocotyl-radicle ratio of soybeans and mung beans and promote the growth of sprouts. To be specific, seed germination rate of soybeans reached the highest in 0.05 mg/L natural brassinolide treatment, which was im- proved by 25.0 percentage points compared with the control group, and the sprout length was improved by 4.33 em; 0.30 mg/L natural brassinolide exhibited the most significant promotion effect on seed germination and sprout growth of mung beans, seed germination rate was improved by 18.3 percentage points and sprout length was improved by 7.29 cm; hypocotyl-radiele ratio of soybean and mung bean sprouts reached the highest （2.96, 1.43 ） in 0.05 mg/L natural brassinolide treatment.

Plant Regeneration from In Vitro Cultured Hypocotyl Explants of Euonymus japonicus Cu zhi

Adventitious shoots were successfully regenerated from hypocotyl explants of in vitro cultures of Euonymus japonicus Cu zhi. Hypocotyl slices were cultured on Murashige and Skoog （MS） and B5 basal medium supplemented with varied concentration of different plant growth-regulators, e.g., α-naphthaleneacetic acid （NAA） and indole-3-butyric acid （IBA） in combination with 6-benzylaminopurine （6-BA） and kinetin. The study showed that shoots could be directly regenerated from hypocotyl explants without the intervening callus phase; MS medium was more suitable for adventitious shoots regeneration. The ability of hypocotyls segments to produce shoots varied depending upon their position on the seedlings. The highest regeneration rate was obtained with hypocotyl segments near to the cotyledon cultured on MS basal medium supplemented with 1.5 mg L^-1 6-BA and 0.05 mg L^-1 NAA （63.64%）. The regenerated shoots were readily elongated on the same medium as used for multiplication and rooted on half-strength MS basal medium supplemented with 1.0 mg L^-1 IBA and 100 mg L^-1 activated carbon. After being transferred to greenhouse conditions, 96% of the plantlets were successfully acclimatized. This regeneration system is applied for genetic transformation now.

Gibberellin A_(3) Treatment Breaks Hypocotyl Dormancy of Paeonia anomala subsp.veitchii Seeds

We studied the roles of different concentrations of gibberellin A_(3)(GA_(3))in breaking hypocotyl dormancy of Paeonia anomala subsp.veitchii,aiming to provide a theoretical basis for the efficient elimination of seed dormancy.The seeds of P.anomala subsp.veitchii were treated with different concentrations of GA_(3)(0,100,200,300,and 400 mg/L)under dark conditions at 16℃,and then the rooting indicators including rooting rate,rooting vigor,rooting index and mold rot rate were determined.The treatment with 400 mg/L GA_(3) for 4 h showed the best performance in breaking the hypocotyl dormancy.Specifically,the treatment shortened the rooting time to 12 d,increased the rooting rate,rooting vigor,and rooting index to 84.00%±5.29%,80.67%±5.03%,and 2.25±0.16 respectively,and decreased the mold rot rate to 0.67%±1.16%.To sum up,there were germination inhibitors in the seeds of P.anomala subsp.veitchii,while the addition of exogenous hormone GA_(3) can release the inhibitory effect on the hypocotyl,which indicated that GA_(3) can promote the breaking of hypocotyl dormancy.

Cortical Microtubule Reorientation and Its Relation to Cell Surface Texture of Epidermal Cells of Arabidopsis Thaliana Hypocotyls under Simulated Microgravity Conditions

Gravitropic curvature growth of Arabidopsis hypocotyls mainly occurred in the rapid growing Elongation Zone(EZI),not in the slow-growing Elongation Zone(EZII).By examining reorientation of Microtubules(MT)and phenotype of the cell wall in the EZI and the EZII of Arabidopsis hypocotyls under normal gravitational condition,it is found that MTs in the rapid growing epidermal cells were mainly in the transverse direction,while those in the non-growing epidermal cells were in the longitudinal directions.However,this difference in cortical MT arrays between the EZI and EZII cells disappeared when the seedlings were exposed to the simulated microgravity condition on a horizontal clinostat.Field emission scanning electron microscopy revealed that the surface texture of epidermal cells,like the direction of the MT,in the EZI and the EZII also became similar when exposed to the simulated microgravity condition.This result indicated that simulate microgravity could modify the potential differentiation between the EZI and the EZII by affecting the orientation of cortical MT in the epidermal cells.

RACK1A promotes hypocotyl elongation by scaffolding light signaling components in Arabidopsis

Plants deploy versatile scaffold proteins to intricately modulate complex cell signaling.Among these,RACK1A(Receptors for Activated C Kinase 1A)stands out as a multifaceted scaffold protein functioning as a central integrative hub for diverse signaling pathways.However,the precise mechanisms by which RACK1A orchestrates signal transduction to optimize seedling development remain largely unclear.Here,we demonstrate that RACK1A facilitates hypocotyl elongation by functioning as a flexible platform that connects multiple key components of light signaling pathways.RACK1A interacts with PHYTOCHROME INTERACTING FACTOR(PIF)3,enhances PIF3 binding to the promoter of BBX11 and down-regulates its transcription.Furthermore,RACK1A associates with ELONGATED HYPOCOTYL 5(HY5)to repress HY5 biochemical activity toward target genes,ultimately contributing to hypocotyl elongation.In darkness,RACK1A is targeted by CONSTITUTIVELY PHOTOMORPHOGENIC(COP)1 upon phosphorylation and subjected to COP1-mediated degradation via the 26 S proteasome system.Our findings provide new insights into how plants utilize scaffold proteins to regulate hypocotyl elongation,ensuring proper skoto-and photo-morphogenic development.

Light-stabilized GIL1 suppresses PIN3 activity to inhibit hypocotyl gravitropism

Light and gravity coordinately regulate the directional growth of plants.Arabidopsis Gravitropic in the Light 1(GIL1)inhibits the negative gravitropism of hypocotyls in red and far-red light,but the underlying molecular mechanisms remain elusive.Our study found that GIL1 is a plasma membrane-localized protein.In endodermal cells of the upper part of hypocotyls,GIL1 controls the negative gravitropism of hypocotyls.GIL1 directly interacts with PIN3 and inhibits the auxin transport activity of PIN3.Mutation of PIN3 suppresses the abnormal gravitropic response of gil1 mutant.The GIL1 protein is unstable in darkness but it is stabilized by red and far-red light.Together,our data suggest that light-stabilized GIL1 inhibits the negative gravitropism of hypocotyls by suppressing the activity of the auxin transporter PIN3,thereby enhancing the emergence of young seedlings from the soil.

The PRR-EC complex and SWR1 chromatin remodeling complex function cooperatively to repress nighttime hypocotyl elongation by modulating PIF4 expression in Arabidopsis

The circadian clock entrained by environmental light-dark cycles enables plants to fine-tune diurnal growth and developmental responses.Here,we show that physical interactions among evening clock components,including PSEUDO-RESPONSE REGULATOR 5(PRR5),TIMING OF CAB EXPRESSION 1(TOC1),and the Evening Complex(EC)component EARLY FLOWERING 3(ELF3),define a diurnal repressive chromatin structure specifically at the PHYTOCHROME-INTERACTING FACTOR 4(PIF4)locus in Arabidopsis.These three clock components act interdependently as well as independently to repress nighttime hypocotyl elongation,as hypocotyl elongation rate dramatically increased specifically at nighttime in the prr5-1 toc1-21 elf3-1 mutant,concomitantly with a substantial increase in PIF4 expression.Transcriptional repression of PIF4 by ELF3,PRR5,and TOC1 is mediated by the SWI2/SNF2-RELATED(SWR1)chromatin remodeling complex,which incorporates histone H2A.Z at thePIF4 locus,facilitating robust epigenetic suppression ofPIF4 during the evening.Overall,these findings demonstrate that the PRR-EC-SWR1 complex represses hypocotyl elongation at night through a distinctive chromatin domain covering PIF4 chromatin.

FERONIA Is a Key Modulator of Brassinosteroid and Ethylene Responsiveness in Arabidopsis Hypocotyls

Ethylene signaling is a complex pathway that has been intensively analyzed partly due to its importance to the manifestation of horticultural phenomena, including fruit ripening and tissue senescence. In order to further our un- derstanding of how this pathway is regulated, a screen for Arabidopsis mutants with increased ethylene response was conducted. From this, a mutant was identified as having a dark-grown hypocotyl that is indistinguishable from Col-0 wt in the presence of the ethylene perception inhibitor AgNO3, yet has extreme responsiveness to even low levels of ethylene. Map-based cloning of the mutation revealed a T-DNA insertion in the coding sequence of the receptor-like kinase FERONIA, which is required for normal pollen tube reception and cell elongation in a currently unknown capacity. In contrast to a previous report, analysis of our feronia knockout mutant shows it also has altered responsiveness to brassinosteroids, with etiolated fer-2 seedlings being partially brassinosteroid insensitive with regard to promotion of hypocotyl elonga- tion. Our results indicate that FERONIA-dependent brassinosteroid response serves to antagonize the effect of ethylene on hypocotyl growth of etiolated seedlings, with loss of proper brassinosteroid signaling disrupting this balance and leading to a greater impact of ethylene on hypocotyl shortening.

Strigolactone-Regulated Hypocotyl Elongation Is Dependent on Cryptochrome and Phytochrome Signaling Pathways in Arabidopsis

Seedling development including hypocotyl elongation is a critical phase in the plant life cycle. Light regula- tion of hypocotyl elongation is primarily mediated through the blue light photoreceptor cryptochrome and red/far-red light photoreceptor phytochrome signaling pathways, comprising regulators including COP1, HY5, and phytochrome- interacting factors （PIFs）. The novel phytohormones, strigolactones, also participate in regulating hypocotyl growth. However, how strigolactone coordinates with light and photoreceptors in the regulation of hypocotyl elongation is largely unclear. Here, we demonstrate that strigolactone inhibition of hypocotyl elongation is dependent on cryp- tochrome and phytochrome signaling pathways. The photoreceptor mutants cry1 cry2, phyA, and phyB are hyposensi- tive to strigolactone analog GR24 under the respective monochromatic light conditions, while cop1 and pifl pif3 pif4 pif5 （pifq） quadruple mutants are hypersensitive to GR24 in darkness. Genetic studies indicate that the enhanced respon- siveness of cop1 to GR24 is dependent on HY5 and MAX2, while that of pifq is independent of HY5. Further studies demonstrate that GR24 constitutively up-regulates HY5 expression in the dark and light, whereas GR24-promoted HY5 protein accumulation is light- and cryptochrome and phytochrome photoreceptor-dependent. These results suggest that the light dependency of strigolactone regulation of hypocotyl elongation is likely mediated through MAX2-dependent promotion of HY5 expression, light-dependent accumulation of HY5, and PIF-regulated components.

Arabidopsis NF-YCs Mediate the Light-Controlled Hypocotyl Elongation via Modulating Histone Acetylation

Light is a crucial environmental signal that promotes photomorphogenesis, the developmental process with a series of light-dependent alterations for plants to adapt various external challenges. Chromatin modification has been proposed to be involved in such light-mediated growth, but the underlying mecha- nism is still elusive. In this study, we identified four Arabidopsis thaliana Nuclear Factor-YC homologs, NF- YCl, NF-YC3, NF-YC4, and NF-YC9 （NF-YCs）, which function redundantly as repressors of light-controlled hypocotyl elongation via histone deacetylation. Obvious etiolation phenotypes are observed in NF-YCs loss-of-function mutant seedlings grown under light conditions, including significant elongated hypocotyls and fewer opened cotyledons. We found that NF-YCs interact with histone deacetylase HDA15 in the light, co-target the promoters of a set of hypocotyl elongation-related genes, and modulate the levels of histone H4 acetylation on the associated chromatins, thus repressing gene expression. In contrast, NF-YC-HDA15 complex is dismissed from the target genes in the dark, resulting in increased level of H4 acetylation and consequent etiolated growth. Further analyses revealed that transcriptional repression activity of NF-YCs on the light-controlled hypocotyl elongation partially depends on the deacetylation activity of HDA15, and loss of HDA15 function could rescue the short-hypocotyl phenotype of NF-YCs overexpression plants. Taken together, our results indicate that NF-YCl, NF-YC3, NF-YC4, and NF-YC9 function as tran- scriptional co-repressors by interacting with HDA15 to inhibit hypocotyl elongation in photomorphogen- esis during the early seedling stage. Our findings highlight that NF-YCs can modulate plant development in response to environmental cues via epigenetic regulation.

The G-Protein β Subunit AGB1 Promotes Hypocotyl Elongation through Inhibiting Transcription Activation Function of BBX21 in Arabidopsis

Hypocotyl development in Arabidopsis thaliana is regulated by light and endogenous hormonal cues, mak- ing it an ideal model to study the interplay between light and endogenous growth regulators. BBX21, a B-box （BBX）-Iike zinc-finger transcription factor, integrates light and abscisic acid signals to regulate hypocotyl elongation in Arabidopsis. Heterotrimeric G-proteins are pivotal regulators of plant development. The short hypocotyl phenotype of the G-protein I^-subunit （AGB1） mutant （agbl-2） has been previously identified, but the precise role of AGB1 in hypocotyl elongation remains enigmatic. Here, we show that AGB1 directly interacts with BBX21, and the short hypocotyl phenotype of agbl-2 is partially suppressed in agb1-2bbx21-1 double mutant. BBX21 functions in the downstream of AGB1 and overexpression of BBX21 in agbl-2 causes a more pronounced reduction in hypocotyl length, indicating that AGB1 plays an oppositional role in relation to BBX21 during hypocotyl development. Furthermore, we demonstrate that the C-terminal region of BBX21 is important for both its intracellular localization and its transcriptional activation activity that is inhibited by interaction with AGB1. ChiP assays showed that BBX21 specifically associates with its own promoter and with those of BBX22, HY5, and GA2oxl. which is not altered in agbl-2. These data suggest that the AGB1-BBX21 interaction only affects the transcrip- tional activation activity of BBX21 but has no effect on its DNA binding ability. Taken together, our data demonstrate that AGB1 positively promotes hypocotyl elongation through repressing BBX21 activity.

Clathrin-Mediated Auxin Efflux and Maxima Regulate Hypocotyl Hook Formation and Light- Stimulated Hook Opening in Arabidopsis

The establishment of auxin maxima by PIN-FORMED 3 （PIN3）- and AUXIN RESISTANT l/LIKE AUX1 （LAX） 3 （AUX1/LAX3）-mediated auxin transport is essential for hook formation in Arabidopsis hypocotyls. Until now, however, the underlying regulatory mechanism has remained poorly understood. Here, we show that loss of function of clathrin light chain CLC2 and CLC3 genes enhanced auxin maxima and thereby hook curvature, alleviated the inhibitory effect of auxin overproduction on auxin maxima and hook curva- ture, and delayed blue light-stimulated auxin maxima reduction and hook opening. Moreover, pharmaco- logical experiments revealed that auxin maxima formation and hook curvature in clc2 clc3 were sensitive to auxin efflux inhibitors 1-naphthylphthalamic acid and 2,3,5-triiodobenzoic acid but not to the auxin influx inhibitor 1-naphthoxyacetic acid. Live-cell imaging analysis further uncovered that loss of CLC2 and CLC3 function impaired PIN3 endocytosis and promoted its lateralization in the cortical cells but did not affect AUX1 localization. Taken together, these results suggest that clathrin regulates auxin maxima and thereby hook formation through modulating PIN3 localization and auxin efflux, providing a novel mechanism that integrates developmental signals and environmental cues to regulate plant skotomorphogenesis and photomorphogenesis.

HOS1 Facilitates the Phytochrome B-Mediated Inhibition of PIF4 Function during Hypocotyl Growth in Arabidopsis

Upon exposure to light, developing seedlings undergo photomorphogenesis, as illustrated by inhibition of hypocotyl elongation, cotyledon opening, and leaf greening. During hypocotyl photomorphogenesis, light signals are sensed by multiple photoreceptors, among which the red/far-red light-sensing phytochromes have been extensively studied. However, it is not fully understood how the phytochromes modulate hypo- cotyl growth. Here, we demonstrated that HIGH EXPRESSION OF OSMOTICALLY RESPONSIVE GENES 1 （HOS1）, which is known to either act as E3 ubiquitin ligase or affect chromatin organization, inhibits the transcriptional activation activity of PHYTOCHROME INTERACTING FACTOR 4 （PIF4）, a key transcrip- tion factor that promotes hypocotyl growth. Consistent with the negative regulatory role of HOSl in hypo- cotyl growth, HOSl-defective mutants exhibited elongated hypocotyls in the light. Notably, phyB induces HOS1 activity in inhibiting PIF4 function. Taken together, these observations provide a molecular basis for the phyB-mediated suppression of hypocotyl growth in Arabidopsis.

The Arabidopsis COP9 SIGNALOSOME INTERACTING F-BOX KELCH 1 Protein Forms an SCF Ubiquitin Ligase and Regulates Hypocotyl Elongation

The regulation of protein turnover by the ubiquitin proteasome system （UPS） is a major posttranslational mechanism in eukaryotes. One of the key components of the UPS, the COP9 signalosome （CSN）, regulates ＇cullin-ring＇ E3 ubiquitin ligases. In plants, CSN participates in diverse cellular and developmental processes, ranging from light signaling to cell cycle control. In this work, we isolated a new plant-specific CSN-interacting F-box protein, which we denominated CFK1 （COP9 INTERACTING F-BOX KELCH 1）. We show that, in Arabidopsis thaliana, CFK1 is a component of a functional ubiquitin ligase complex. We also show that CFK1 stability is regulated by CSN and by proteasome-dependent proteoly- sis, and that light induces accumulation of the CFK1 transcript in the hypocotyl. Analysis of CFK1 knockdown, mutant, and overexpressing seedlings indicates that CFK1 promotes hypocotyl elongation by increasing cell size. Reduction of CSN levels enhances the short hypocotyl phenotype of CFKl-depleted seedlings, while complete loss of CSN activity sup- presses the Iong-hypocotyl phenotype of CFKl-overexpressing seedlings. We propose that CFK1 （and its regulation by CSN） is a novel component of the cellular mechanisms controlling hypocotyl elongation.

REVEILLE 7 inhibits the expression of the circadian clock gene EARLY FLOWERING 4 to fine-tune hypocotyl growth in response to warm temperatures

The circadian clock maintains the daily rhythms of plant growth and anticipates predictable ambient temperature cycles.The evening complex(EC),comprising EARLY FLOWERING 3(ELF3),ELF4,and LUX ARRHYTHMO,plays an essential role in suppressing thermoresponsive hypocotyl growth by negatively regulating PHYTOCHROME INTERACTING FACTOR 4(PIF4)activity and its downstream targets in Arabidopsis thaliana.However,how EC activity is attenuated by warm temperatures remains unclear.Here,we demonstrate that warm temperature-induced REVEILLE7(RVE7)fine-tunes thermoresponsive growth in Arabidopsis by repressing ELF4 expression.RVE7 transcript and RVE7 protein levels increased in response to warm temperatures.Under warm temperature conditions,an rve7 loss-of-function mutant had shorter hypocotyls,while overexpressing RVE7 promoted hypocotyl elongation.PIF4 accumulation and downstream transcriptional effects were reduced in the rve7 mutant but enhanced in RVE7 overexpression plants under warm conditions.RVE7 associates with the Evening Element in the ELF4 promoter and directly represses its transcription.ELF4 is epistatic to RVE7,and overexpressing ELF4 suppressed the phenotype of the RVE7 overexpression line under warm temperature conditions.Together,our results identify RVE7 as an important regulator of thermoresponsive growth that functions(in part)by controlling ELF4 transcription,highlighting the importance of ELF4 for thermomorphogenesis in plants.

The E3 ligase XBAT35 mediates thermoresponsive hypocotyl growth by targeting ELF3 for degradation in Arabidopsis

Plants are capable of coordination of their growth and development with ambient temperatures.EARLY FLOWERING3(ELF3), an essential component of the plant circadian clock, is also involved in ambient temperature sensing, as well as in inhibiting the expression and protein activity of the thermoresponsive regulator phytochrome interacting factor4(PIF4). The ELF3 activity is subjected to attenuation in response to warm temperature;however,how the protein level of ELF3 is regulated at warm temperature remains less understood. Here, we report that the E3 ligase XB3 ORTHOLOG 5 IN ARABIDOPSIS THALIANA, XBAT35, mediates ELF3 degradation. XBAT35 interacts with ELF3 and ubiquitinates ELF3. Loss-of-function mutation of XBAT35 increases the protein level of ELF3 and confers a short-hypocotyl phenotype under warm temperature conditions. Thus, our findings establish that XBAT35 mediates ELF3 degradation to lift the inhibition of ELF3 on PIF4 for promoting thermoresponsive hypocotyl growth in plants.

The DELLA-ABI4-HY5 module integrates light and gibberellin signals to regulate hypocotyl elongation

Plant growth is coordinately controlled by various environmental and hormonal signals,of which light and gibberellin(GA)signals are two critical factors with opposite effects on hypocotyl elongation.Although interactions between the light and GA signaling pathways have been studied extensively,the detailed regulatory mechanism of their direct crosstalk in hypocotyl elongation remains to be fully clarified.Previously,we reported that ABA INSENSITIVE 4(ABI4)controls hypocotyl elongation through its regulation of cellelongation-related genes,but whether it is also involved in GA signaling to promote hypocotyl elongation is unknown.In this study,we showthat promotion of hypocotyl elongation by GA is dependent on ABI4 activation.DELLAs interact directly with ABI4 and inhibit its DNA-binding activity.In turn,ABI4 combined with ELONGATED HYPOCOTYL 5(HY5),a key positive factor in light signaling,feedback regulates the expression of the GA2ox GA catabolism genes and thus modulates GA levels.Taken together,our results suggest that the DELLA-ABI4-HY5 module may serve as a molecular link that integrates GA and light signals to control hypocotyl elongation.