An Evolutionary Strategy for Spawning Habitat Selection by Pseudopungtungia tenuicorpa.

In this study, we hypothesized that the limited geographical distribution of Pseudopungtungia tenuicorpa in South Korea is closely related to the local abundance of their preferred spawning habitats. Eggs of P. tenuicorpa were most frequently observed in cracks smaller than 5 cm, followed by the nests of other fish, such as Coreoperca herzi. Although eggs of P. tenuicorpa were also found in cracks larger than 6 cm, most of them were consumed or damaged by other fish. However, the number of P. tenuicorpa eggs spawned in cracks smaller than 5 cm remained similar to the number of eggs at the time of initial spawning, even after 9 days. The strong preference of P. tenuicorpa for cracks smaller than 5 cm was found to influence the geographical distribution of P. tenuicorpa in South Korea. In the upper reaches of the Han River, the ratio of cracks smaller than 5 cm is higher than that in the Geum River and in the Nakdong River, which contributes to the stable population growth of P. tenuicorpa there. In the future, management strategies to maintain/restore cracks smaller than 5 cm should be reviewed for the preservation of this endemic fish in South Korea.

Behavioral Avoidance Response of Daphnia to Fungal Infection Caused by Metschnikowia Species in a Temperate Reservoir.

Morphological or behavioral defense mechanisms are important evolutionary strategies for the survival of prey. Studies have focused on predation and competition, but infection has been overlooked, despite being a determining factor of distribution and species diversity of prey. We hypothesized that the winter migration of Daphnia pulicaria is a community defense strategy to avoid fungal infection. To test this hypothesis, environmental variables and the Cladocera community, including D. pulicaria, were monitored in three study sections of the Anri Reservoir in the Republic of Korea during September 2010-August 2015. During three winter seasons, the density of infected D. pulicaria increased in all study sections, and they migrated from the central to the littoral area. Most of the infected individuals had dormant eggs in sexually reproducing mothers. However, when the proportion of non-infected individuals was higher than that of infected individuals, winter migration was not observed. Additional microcosm experiments showed that dormant eggs of D. pulicaria obtained from ice crystals in the littoral area had lower hatching and infection rates than those obtained from mothers moving from other zones. Therefore, the migration of D. pulicaria during winter is an active response to avoid intergenerational fungal infection.

Deconvoluting signals downstream of growth and immune receptor kinases by phosphocodes of the BSU1 family phosphatases.

Hundreds of leucine-rich repeat receptor kinases (LRR-RKs) have evolved to control diverse processes of growth, development and immunity in plants, but the mechanisms that link LRR-RKs to distinct cellular responses are not understood. Here we show that two LRR-RKs, the brassinosteroid hormone receptor BRASSINOSTEROID INSENSITIVE 1 (BRI1) and the flagellin receptor FLAGELLIN SENSING 2 (FLS2), regulate downstream glycogen synthase kinase 3 (GSK3) and mitogen-activated protein (MAP) kinases, respectively, through phosphocoding of the BRI1-SUPPRESSOR1 (BSU1) phosphatase. BSU1 was previously identified as a component that inactivates GSK3s in the BRI1 pathway. We surprisingly found that the loss of the BSU1 family phosphatases activates effector-triggered immunity and impairs flagellin-triggered MAP kinase activation and immunity. The flagellin-activated BOTRYTIS-INDUCED KINASE 1 (BIK1) phosphorylates BSU1 at serine 251. Mutation of serine 251 reduces BSU1's ability to mediate flagellin-induced MAP kinase activation and immunity, but not its abilities to suppress effector-triggered immunity and interact with GSK3, which is enhanced through the phosphorylation of BSU1 at serine 764 upon brassinosteroid signalling. These results demonstrate that BSU1 plays an essential role in immunity and transduces brassinosteroid-BRI1 and flagellin-FLS2 signals using different phosphorylation sites. Our study illustrates that phosphocoding in shared downstream components provides signalling specificities for diverse plant receptor kinases.

Brassinosteroids enhance salicylic acid-mediated immune responses by inhibiting BIN2 phosphorylation of clade I TGA transcription factors in Arabidopsis.

Salicylic acid (SA) plays an important role in plant immune response, including resistance to pathogens and systemic acquired resistance. Two major components, NONEXPRESSOR OF PATHOGENESIS-RELATED GENES (NPRs) and TGACG motif-binding transcription factors (TGAs), are known to mediate SA signaling, which might also be orchestrated by other hormonal and environmental changes. Nevertheless, the molecular and functional interactions between SA signaling components and other cellular signaling pathways remain poorly understood. Here we showed that the steroid plant hormone brassinosteroid (BR) promotes SA responses by inactivating BR-INSENSITIVE 2 (BIN2), which inhibits the redox-sensitive clade I TGAs in Arabidopsis. We found that both BR and the BIN2 inhibitor bikinin synergistically increase SA-mediated physiological responses, such as resistance to Pst DC3000. Our genetic and biochemical analyses indicated that BIN2 functionally interacts with TGA1 and TGA4, but not with other TGAs. We further demonstrated that BIN2 phosphorylates Ser-202 of TGA4, resulting in the suppression of the redox-dependent interaction between TGA4 and NPR1 as well as destabilization of TGA4. Consistently, transgenic Arabidopsis overexpressing TGA4-YFP with a S202A mutation displayed enhanced SA responses compared to the wild-type TGA4-YFP plants. Taken together, these results suggest a novel crosstalk mechanism by which BR signaling coordinates the SA responses mediated by redox-sensitive clade I TGAs.

Hydrological Characteristics and Trophic Status as Dominant Drivers of Rotifer Community Composition in Artificially Created Riverine Wetlands.

Hydrological characteristics of freshwater ecosystems are powerful determinants of the distribution of biological communities and changes in environmental factors. This study identified relationships between the wetland environment, rotifer community, and hydrological factors for 48 wetlands, to determine their impact on wetland conservation and management. Different hydrological factors produced different wetland environments, which influenced the rotifer community composition. The wetlands with "poor" "in/outflow function" and "shore stability" levels showed high conductivity, turbidity, depth, and concentrations of total nitrogen, total phosphorus, and chlorophyll a. In contrast, the dissolved oxygen levels and velocity were the highest in wetlands with "good" in/outflow function and shore stability variables. The nutritional status of each wetland affected the composition of the rotifer community. Some genera (Keratella, Brachionus, Anuraeopsis, Trichocerca, and Philodina) were found in wetlands with high concentrations of total nitrogen, total phosphorus, and chlorophyll a, and high turbidity and depth. In contrast, Ascomorpha and Ploesoma were found in wetlands with high dissolved oxygen levels and flow velocity. High densities of Lepadella, Lecane, and Testudinella were found in wetlands completely covered by macrophytes. The rotifer community distribution can be used to understand the trophic, current functional, and environmental status of wetlands.

Inhibition of 4-HYDROXYPHENYLPYRUVATE DIOXYGENASE expression by brassinosteroid reduces carotenoid accumulation in Arabidopsis.

Unlike the indispensable function of the steroid hormone brassinosteroid (BR) in regulating plant growth and development, the metabolism of secondary metabolites regulated by BR is not well known. Here we show that BR reduces carotenoid accumulation in Arabidopsis seedlings. BR-deficient or BR-insensitive mutants accumulated higher content of carotenoids than wild-type plants, whereas BR treatment reduced carotenoid content. We demonstrated that BR transcriptionally suppresses 4-HYDROXYPHENYLPYRUVATE DIOXYGENASE (HPPD) expression involved in carotenogenesis via plastoquinone production. We found that the expression of HPPD displays an oscillation pattern that is expressed more strongly in dark than in light conditions. Moreover, BR appeared to inhibit HPPD expression more strongly in darkness than in light, leading to suppression of a diurnal oscillation of HPPD expression. BR-responsive transcription factor BRASSINAZOLE RESISTANT 1 (BZR1) directly bound to the promoter of HPPD, and HPPD suppression by BR was increased in the bzr1-1D gain-of-function mutation. Interestingly, dark-induced HPPD expression did not cause carotenoid accumulation, due to down-regulation of other carotenoid biosynthetic genes in the dark. Our results suggest that BR regulates different physiological responses in dark and light through inhibition of HPPD expression.

Invasion and Dispersion of the Exotic Species Procambarus clarkii (Decapoda Cambaridae) in Yeongsan River Basin, South Korea.

The introduction of exotic species negatively affects the distribution and interactions within local biological communities in an ecosystem and can threaten ecosystem health. This study aimed to provide the basic data required to manage P. clarkii in the Yeongsan River basin. We identified the dispersion pattern and evaluated the ecosystem risk of this newly introduced species. The distribution survey investigated Procambarus clarkii populations at 25 sites in the Yeongsan River basin over a four-year period. The initial introduction occurred in Jiseok Stream. The larvae of P. clarkii were most abundant in areas with a dense aquatic plant cover, whereas adults preferred silt/clay areas. The alterations in the water flow by the river refurbishment project (carried out in 2012) increased their preferred habitats and contributed to P. clarkii dispersion. However, stable isotope analysis showed that the dispersion has had little effect on the freshwater ecosystem. The interrelationship between P. clarkii (i.e., larvae and adults) and other biological communities has been limited. Although the rapid dispersion by P. clarkii in the Yeongsan River basin has not impacted the freshwater ecosystem, further ecological information is required on how to manage P. clarkii beyond this early stage of invasion.

Changes in Antibiotic-Resistance Genes Induced by the Grazing Effect in Three Cladoceran Species.

The acquisition of Antibiotic-Resistance Genes (ARGs) by natural bacteria caused by antibiotic abuse is causing serious problems for human and animal welfare. Here, we evaluated the influence of three cladoceran species on Antibiotic-Resistant Bacteria (ARB) and tetracycline-resistance gene (tet(A)) copies, and discussed the effect of these biological interactions on the distribution and diffusion of ARGs in freshwater ecosystems. Bacterial community and tet(A) abundances in water samples collected from wetlands were strongly influenced by cladoceran presence. The presence of Daphnia obtusa dramatically decreased ARB and tet(A) abundance compared to that with other cladoceran species (Chydorus sphaericus and Simocephalus vetulus). Interestingly, we found a high abundance of Flavobacteriales in the microbiomes of cladoceran species. Considering that Flavobacteriales species are potential carriers of the tet(A) gene, their adsorption and assimilation with cladocerans could significantly impact the reduction of tet(A) in water. Field surveys also showed that tet(A) abundance could be low if the dominance of D. obtusa in each wetland was high. This study highlighted the need for ecological interactions and a broad range of niches in the food web when discussing the fate of ARGs in freshwater ecosystems.

The Use of Winter Water Temperature and Food Composition by the Copepod Cyclops vicinus (Uljanin, 1875) to Provide a Temporal Refuge from Fish Predation.

Frequent predation induces various defense strategies in prey, including morphological changes or migration patterns in zooplankton. We hypothesized that the winter dominance of Cyclops vicinus in the Upo Wetlands, South Korea, is an evolved temporal defense mechanism to avoid fish predation. Long-term data (2014-2019) showed that fish consumed the most cyclopoid copepods from spring to autumn. Lepomis macrochirus preferentially consumed C. vicinus; thus, C. vicinus density was lower from spring to autumn. However, C. vicinus was abundant in winter when fish consumed fewer copepods. Nauplii density began to increase in late autumn (October-November), and their population growth was fueled through consumption of Cyclotella sp. and Rhodomonas sp. Culture experiments showed that Cyclotella sp. contributed more to the growth stage (copepodite or subadult) after nauplii than Rhodomonas sp. C. vicinus density was lower in the winters of 2013 and 2016 when the densities of these phytoplankton prey species were lower. In summary, although winter conditions were suitable for copepod survival and population growth, C. vicinus relied heavily on the diversity and species composition of its food sources. The winter dominance of C. vicinus could increase regional biodiversity and contribute significantly to the stability of the freshwater food web.

Endogenous level of abscisic acid down-regulated by brassinosteroids signaling via BZR1 to control the growth of Arabidopsis thaliana.

The increased level of endogenous abscisic acid (ABA) in brassinosteroid (BR)-deficient mutants, such as det2 and cyp85a1 × cyp85a2, suggests that ABA synthesis is inhibited by endogenous BRs in Arabidopsis thaliana. Expression of the ABA biosynthesis gene ABA-deficient 2 (ABA2) was negatively regulated by exogenously applied BR but up-regulated by the application of brassinazole and in det2 and cyp85a1 × cyp85a2. In addition, ABA2 expression decreased in bzr1-1D, showing that ABA biosynthesis is inhibited by BR signaling via BZR1, intermediated by ABA2, in Arabidopsis. Four cis-element sequences (E-boxes 1-4) in the putative promoter region of ABA2 were identified as BZR1 binding sites. The electrophoretic mobility shift assay and chromatin immune precipitation analysis demonstrated that BZR1 directly binds to overlapped E-boxes (E-box 3/4) in the promoter region of ABA2. The level of endogenous ABA was decreased in bzr1-1D compared to wild-type, indicating that binding of BZR1 to the ABA2 promoter inhibits ABA synthesis in Arabidopsis. Compared to wild-type, aba2-1 exhibited severely reduced growth and development. The abnormalities in aba2-1 were rescued by the application of ABA, suggesting that ABA2 expression and ABA synthesis are necessary for the normal growth and development of A. thaliana. Finally, bzr1-KO × aba2-1 exhibited inhibitory growth of primary roots compared to bzr1-KO, verifying that ABA2 is a downstream target of BZR1 in the plant. Taken together, the level of endogenous ABA is down-regulated by BR signaling via BZR1, controlling the growth of A. thaliana.

Seed-Specific Expression of Arabidopsis AtCYP85A2 Produces Biologically Active Brassinosteroids Such as Castasterone and Brassinolide to Improve Grain Yield and Quality in Seeds of Brachypodium Distachyon.

Brachypodium distachyon is a monocotyledonous model plant that contains castasterone (CS) but no brassinolide (BL) as the end product of brassinosteroids (BR) biosynthesis, indicating dysfunction of BL synthase, which catalyzes the conversion of CS to BL. To increase BR activity, Arabidopsis cytochrome P450 85A2 (AtCYP85A2) encoding BR 6-oxidase/BL synthase was introduced into B. distachyon with the seed-specific promoters pBSU1, pAt5g10120, and pAt5g54000. RT-PCR analysis and GUS activity revealed that AtCYP85A2 was only expressed in the seeds of the transgenic plants pBSU1-AtCYP85A2::Bd21-3, pAt5g10120-AtCYP85A2::Bd21-3, and pAt5g54000-AtCYP85A2::Bd21-3. The crude enzyme prepared from the seeds of these three transgenic plants catalyzed the conversion of 6-deoxoCS to CS. The transgenic plants showed greater specific enzyme activity than the wild-type plant for the conversion of 6-deoxoCS to CS, indicating enhanced BR 6-oxidase activity in the transgenic plants. The enzyme solution also catalyzed the conversion of CS into BL. Additionally, BL was identified from the seeds of transgenic plants, verifying that seed-specific AtCYP85A2 encodes a functional BL synthase to increase BR activity in the seeds of transgenic Brachypodium. In comparison with wild-type Brachypodium, the transgenic plants showed better growth and development during the vegetative growing stage. The flowers of the transgenic plants were remarkably larger, resulting in increments in the number, size, and height of seeds. The total starch, protein, and lipid contents in transgenic plants were higher than those in wild-type plants, indicating that seed-specific expression of AtCYP85A2 improves both grain yield and quality in B. distachyon.

BES1 negatively regulates the expression of ACC oxidase 2 to control the endogenous level of ethylene in Arabidopsis thaliana.

Quantitative reverse transcription PCR (qRT-PCR) analysis and ProACO2::GUS expression showed that ACO2 was highly expressed in the shoots of Arabidopsis seedlings under light conditions. Exogenously applied aminocyclopropane-1-carboxylic acid (ACC) enhanced the expression of ACO2, whereas Co2+ ions suppressed its expression. In comparison with wild-type seedlings, the ACO2 knockdown mutant aco2-1 produced less ethylene, which resulted in the inhibited growth of Arabidopsis seedlings. Exogenously applied brassinolide reduced the expression of ACO2. ACO2 expression was increased in det2, a brassinosteroid (BR)-deficient mutant; however, it was decreased in bes1-D, a brassinosteroid insensitive 1-EMS-suppressor 1 (BES1)-dominant mutant. In the putative promoter region of ACO2, 11 E-box sequences for BES1 binding but not BR regulatory element sequences for brassinazole-resistant 1 (BZR1) binding were found. Chromatin immunoprecipitation assay showed that BES1 could directly bind to the E-boxes located in the putative promoter region of ACO4. Less ethylene was produced in bes1-D seedlings compared with wild-type seedlings, suggesting that the direct binding of BES1 to the ACO2 promoter may negatively regulate ACO2 expression to control the endogenous level of ethylene in Arabidopsis seedlings.

Habitat Preferences and Trophic Position of Brachydiplax chalybea flavovittata Ris, 1911 (Insecta: Odonata) Larvae in Youngsan River Wetlands of South Korea.

In freshwater ecosystems, habitat heterogeneity supports high invertebrate density and diversity, and it contributes to the introduction and settlement of non-native species. In the present study, we identified the habitat preferences and trophic level of Brachydiplax chalybea flavovittata larvae, which were distributed in four of the 17 wetlands we examined in the Yeongsan River basin, South Korea. Larval density varied across four microhabitat types: open water area, and microhabitats dominated by Myriophyllum aquaticum, Paspalum distichum, and Zizania latifolia. Microhabitats dominated by M. aquaticum had the highest larval density, followed by those dominated by P. distichum. The larvae were more prevalent in silt sediments than in plant debris or sand. Stable isotope analysis showed that B. chalybea flavovittata is likely to consume, as a food source, other species of Odonata larvae. We conclude that successful settlement of B. chalybea flavovittata can be attributed to their habitat preferences. As temperature increases due to climate change, the likelihood of B. chalybea flavovittata spreading throughout South Korea increases. We, therefore, recommend continued monitoring of the spread and ecological impacts of B. chalybea flavovittata.

Brassinosteroids signaling via BZR1 down-regulates expression of ACC oxidase 4 to control growth of Arabidopsis thaliana seedlings.

ProACO4-GUS expression and RT-PCR analysis revealed that ACO4 is predominantly expressed in shoots of Arabidopsis seedlings under light conditions. ACO4-overexpressed mutant 35S-ACO4 produced more ethylene relative to the wild-type, which resulted in reduced growth of Arabidopsis seedlings. The abnormal growth of seedlings recurred after the application of Co2+ ions, suggesting that ACO4 is a functional ACO necessary to regulate the growth and development of Arabidopsis seedlings. Exogenously-applied brassinosteroids (BRs) inhibited the expression of ACO4, and an enhanced ACO4 expression was found in det2, a BR-deficient mutant. Additionally, expression of ACO4 was decreased in bzr1-D (a BZR1-dominant mutant), implying that BR signaling negatively regulates ACO4 expression via BZR1 in Arabidopsis. In the intergenic region of ACO4, four E-boxes and a BR regulatory element (BRRE) are found. Electrophoretic mobility shift and chromatin immunoprecipitation assays showed that BZR1 binds directly to the BRRE in the putative promoter region of ACO4. By binding of BZR1 to BRRE, less ethylene was produced, which seems to regulate the growth and development of Arabidopsis seedlings.

Establishment of Biosynthetic Pathways To Generate Castasterone as the Biologically Active Brassinosteroid in Brachypodium distachyon.

Gas chromatography-mass spectrometry (GC-MS) analysis revealed that castasterone and its biosynthetic precursors are found in Brachypodium distachyon. In vitro conversion experiments with crude enzyme solutions prepared from B. distachyon demonstrated the presence of the following biosynthetic sequences: campesterol → campesta-4-en-3-one → campesta-3-one → campestanol → 6-deoxocathasterone → 6-deoxoteasterone → teasterone ↔ 3-dehydroteasterone ↔ typhasterol → castasterone. campesterol → 22-hydroxycampesterol → 22-hydroxy-campesta-4-en-3-one → 22-hydroxy-campesta-3-one → 6-deoxo-3-dehydroteasterone → 3-dehydroteasterone. 6-deoxoteasterone ↔ 6-deoxo-3-dehydroteasterone ↔ 6-deoxotyphasterol → 6-deoxocastasterone → castasterone. This shows that there are campestanol-dependent and campestanol-independent pathway in B. distachyon that synthesize 24-methylated brassinosteroids (BRs). Biochemical analysis of BRs biosynthetic enzymes confirmed that BdDET2, BdCYP90B1, BdCYP90A1, BdCYP90D2, and BdCYP85A1 are orthologous to BR 5α-reductase, BR C-22 hydroxylase, BR C-3 oxidase, BR C-23 hydroxylase, and BR C-6 oxidase, respectively. Brassinolide was not identified in B. distachyon. Additionally, B. distachyon crude enzyme solutions could not catalyze the conversion of castasterone to brassinolide, and the gene encoding an ortholog of CYP85A2 (a brassinolide synthase) was not found in B. distachyon. These results strongly suggest that the end product for brassinosteroid biosynthesis which controls the growth and development of B. distachyon is not brassinolide but rather castasterone.

BES1 directly binds to the promoter of the ACC oxidase 1 gene to regulate gravitropic response in the roots of Arabidopsis thaliana.

Brassinosteroids (BRs) are known to be endogenous regulators of ethylene production, suggesting that some BR activity in plant growth and development is associated with ethylene. Here, we demonstrated that ethylene production in Arabidopsis thaliana roots is increased by BR signaling via the ethylene biosynthetic gene for ACC oxidase 1 (ACO1). Electrophoretic mobility shift and chromatin immune-precipitation assays showed that the BR transcription factor BES1 directly binds to two E-box sequences located in the intergenic region of ACO1. GUS expression using site mutations of the E-box sequences verified that ACO1 is normally expressed only when BES1 binds to the E-boxes in the putative promoter of ACO1, indicating that this binding is essential for ACO1 expression and the subsequent production of ethylene in A. thaliana roots. BR exogenously applied to A. thaliana roots enhanced the gravitropic response. Additionally, bes1-D exhibited a greater gravitropic response than did the wild-type specimens, proving that BR is a positive regulator of the gravitropic response in A. thaliana roots. The knock-down mutant aco1-1 showed a slightly lower gravitropic response than did the wild-type specimens, while bes1-D X aco1-1 exhibited a lower gravitropic response than did bes1-D. Therefore, ACO1 is a direct downstream target for BR transcription factor BES1, which controls ethylene production for gravitropism in A. thaliana roots.

Plant U-Box40 Mediates Degradation of the Brassinosteroid-Responsive Transcription Factor BZR1 in Arabidopsis Roots.

Brassinosteroid (BR) regulates a wide range of physiological responses through the activation of BRASSINAZOLE RESISTANT1 (BZR1), whose activity is tightly controlled by its phosphorylation status and degradation. Although BZR1 appears to be degraded in distinct ways in response to different hormonal or environmental cues, little is known about how BR signaling regulates its degradation. Here we show that the BR-regulated U-box protein PUB40 mediates the proteasomal degradation of BZR1 in a root-specific manner in Arabidopsis (Arabidopsis thaliana). BZR1 levels were strongly reduced by plant U-box40 (PUB40) overexpression, whereas the pub39 pub40 pub41 mutant accumulated much more BZR1 than wild type in roots. The bzr1-1D gain-of-function mutation reduced the interaction with PUB40, which suppressed PUB40-mediated BZR1 degradation in roots. The cell layer-specific expression of PUB40 in roots helps induce selective BZR1 accumulation in the epidermal layer. Both BR treatment and loss-of-function of PUB40 expanded BZR1 accumulation to most cell layers. In addition, BZR1 accumulation increased the resistance of pub39 pub40 pub41 to low inorganic phosphate availability, as observed in bzr1-1D BRASSINOSTEROID-INSENSITIVE2-induced phosphorylation of PUB40, which mainly occurs in roots, gives rise to BZR1 degradation through enhanced binding of PUB40 to BZR1 and PUB40's stability. Our results suggest a molecular mechanism of root-specific BZR1 degradation regulated by BR signaling.

The complete chloroplast genome of Korean endemic species, Cirsium rhinoceros (H.Lév. & vaniot) Nakai (Asteraceae).

Cirsium rhinoceros (H.Lév. & Vaniot) Nakai has been used a traditional medicine. Complete chloroplast genome of C. rhinoceros is 152,576 bp long and has four subregions: 87,262 bp of large single copy (LSC) and 21,486 bp of small single copy (SSC) regions that are separated by 18,742 bp of inverted repeat (IR) regions including 133 genes (88 protein-coding genes, 8 rRNAs, and 37 tRNAs). The overall GC content of this chloroplast genome is 37.7% and in the LSC, SSC, and IR regions are 36.0%, 31.4%, and 43.8%, respectively. Phylogenetic trees show that Cirsium species are clustered along with their distribution.

Arabidopsis ACC Oxidase 1 Coordinated by Multiple Signals Mediates Ethylene Biosynthesis and Is Involved in Root Development.

Ethylene regulates numerous aspects of plant growth and development. Multiple external and internal factors coordinate ethylene production in plant tissues. Transcriptional and post-translational regulations of ACC synthases (ACSs), which are key enzymes mediating a rate-limiting step in ethylene biosynthesis have been well characterized. However, the regulation and physiological roles of ACC oxidases (ACOs) that catalyze the final step of ethylene biosynthesis are largely unknown in Arabidopsis. Here, we show that Arabidopsis ACO1 exhibits a tissue-specific expression pattern that is regulated by multiple signals, and plays roles in the lateral root development in Arabidopsis. Histochemical analysis of the ACO1 promoter indicated that ACO1 expression was largely modulated by light and plant hormones in a tissue-specific manner. We demonstrated that point mutations in two E-box motifs on the ACO1 promoter reduce the light-regulated expression patterns of ACO1. The aco1-1 mutant showed reduced ethylene production in root tips compared to wild-type. In addition, aco1-1 displayed altered lateral root formation. Our results suggest that Arabidopsis ACO1 integrates various signals into the ethylene biosynthesis that is required for ACO1's intrinsic roles in root physiology.