The growth-defence trade-offs pose a major challenge to breeding high-yield and disease-resistant crops. Aquaporins are membrane channels that facilitate the transport of water and other small compounds, therefore regulating the growth-defence trade-offs. However, the molecular mechanism that governs the function of aquaporins in the trade-offs remains unclear. Here, we report that Triticum aestivum TaPIP1;6 and TaPIP2;10, aquaporins of the plasma membrane intrinsic protein (PIP) family, function as dual substrate channels that concurrently enhance plant growth and resistance to powdery mildew and English grain aphid. In wheat plants growing under normal conditions, TaPIP1;6 and TaPIP2;10 facilitate CO2 transport from the atmosphere into wheat cells and promote photosynthesis, which leads to growth enhancement and grain yield increase. In wheat plants under attack by powdery mildew pathogen or English grain aphid, TaPIP1;6 and TaPIP2;10 function as concurrent H2O2 transport channels, mediating the influx of apoplastic H2O2 into the cytoplasm. In turn, the transported H2O2 activates innate immunity, including the MAPK cascade, callose deposition and defence gene expression, and thereby enhances wheat resistance to powdery mildew and the English grain aphid. In essence, co-overexpression of TaPIP1;6 and TaPIP2;10 exhibits synergistic effects on CO2 and H2O2 transports, further amplifying both yield and resistance traits. Taken together, our results suggest that TaPIP1;6 and TaPIP2;10 function as dual-substrate transporting channels to promote growth by regulating CO2 transport and to enhance resistance against pathogens and insects via H2O2-mediated immune responses. This finding provides crucial genetic targets for breeding crop varieties combining high yield with resistance traits.

Abstract To advance modern crop breeding by de novo domestication, Hui Lin and colleagues recently characterized an immune module in rice subspecies Oryza sativa subsp. indica (OSI) and O. sativa subsp. japonica (OSJ). The module involves rice resistance protein XA48 and the downstream transcription factor OsVOZ1. Carrying an alanine (A) to serine (S) variation, OsVOZ1 functions redundantly with OsVOZ2 in suppressing rice resistance to bacterial blight, a worldwide significant crop disease caused by Xanthomonas oryzae pv. oryzae (Xoo). XA48, identified in an OSI cultivar, recognizes the Xoo effector XopG, confers resistance and susceptibility to XopG-harboring Xoo strains from Northeast and South Asia, respectively, and activates effector-triggered immunity (ETI) by degrading OsVOZ1/2. In resistance spectra, this ETI complements pattern-triggered immunity (PTI) mediated by XA21, a resistance protein previously identified in wild rice. The Xa48–OsVOZ1 module has undergone asymmetric selection: Xa48 is retained in OSI and lost in OSJ, whereas OSI retains both OsVOZ1 A/S alleles and OSJ retains only OsVOZ1 A . Although restoring XA48 to OSJ severely reduces grain yield, integrating XA48/ETI and XA21/PTI into modern OSI and OSJ cultivars rebuilds broad-spectrum resistance without a yield penalty, opening an avenue toward modern crop breeding using immunity sources from wild rice.

Fusarium graminearum is recognized as the pathogen responsible for wheat head blight. It produces deoxynivalenol (DON) during infection, which endangers human health. DON biosynthesis occurs within toxisomes in the endoplasmic reticulum (ER). In eukaryotes, the ER membrane protein complex (EMC) is critical for the ER’s normal operation. However, the specific role of the EMC in F. graminearum remains poorly understood. In this study, six EMC subunits (FgEmc1-6) were identified in F. graminearum, and all of them were localized to the toxisomes. Our results demonstrate that the EMC is indispensable for vegetative growth and asexual and sexual reproduction, which are the fundamental life processes of F. graminearum. Importantly, EMC deletion led to reduced virulence in wheat spikes and petioles. Further investigation revealed that in ΔFgemc1-6, the expression of trichothecene (TRI) genes is decreased, the biosynthesis of lipid droplets (LDs) is diminished, toxisome formation is impaired, and DON production is reduced. Additionally, defects in the formation of the infection cushion were observed in ΔFgemc1-6. In conclusion, the EMC is involved in regulating growth and virulence in F. graminearum. This study enhances our understanding of the EMC functions in F. graminearum and offers valuable insights into potential targets for managing wheat head blight.

TRAnsport Protein Particle (TRAPP) is a conserved multi-subunit tethering complex known to be involved in intracellular movement of proteins. However, its components and molecular functions in filamentous fungi remain poorly characterized. Here, we identify four TRAPPIII-specific subunits (FgTrs85, TRAPPC11, TRAPPC12, and TRAPPC13) in the phytopathogenic fungus Fusarium graminearum. Genetic and functional analyses reveal that FgTrs85 serves as the core subunit, collaborating with auxiliary subunits TRAPPC11, TRAPPC12, and TRAPPC13 to orchestrate fungal perithecium formation, growth, and virulence. TRAPPIII localizes to the phagophore assembly site and promotes autophagosome biogenesis by recruiting FgAtg9 through interactions involving FgTrs85 and TRAPPC13. Notably, TRAPPIII mutants exhibit more severe growth defects than autophagy-deficient strains, suggesting that the roles of TRAPPIII extend beyond autophagy. TRAPPIII regulates ER-to-Golgi and endosome-to-Golgi transport by ensuring the proper localization of secretory regulators (FgSec22, FgRud3, FgSnc1). Moreover, the overexpression of FgRab1-GTP largely suppresses all phenotypic defects associated with perithecium formation, growth, and virulence in TRAPPIII mutants, suggesting the function of TRAPPIII as a guanine nucleotide exchange factor that activates FgRab1. Altogether, our results demonstrate that TRAPPIII coordinates autophagy and intracellular transport to regulate fungal development, growth and virulence in F. graminearum.

Plant proteins that belong to the nonexpressor of pathogenesis-related (NPR) gene family are paralogous receptors of the plant defense hormone salicylic acid and essential regulators of hormone-dependent plant immunity against diseases caused by various pathogens. Previous studies have established NPR1 and NPR3 as a transcriptional activator and a transcriptional repressor, respectively, of defense-gene expression to promote and inhibit broad-spectrum resistance against different strains of pathogens. However, the regulatory mechanism that underlies the opposing roles of NPR1 and NPR3 in defense-gene activation remains unclear. Here, we report that a rice transcript splicing factor, Oryza sativa RNA-binding protein 11 (OsRBP11), promotes alternative splicing of OsNPR3 to modulate the defense function of OsNPR1 in rice plants infected by Xanthomonas oryzae pathovars, which are important bacterial pathogens of rice. We discovered that 11 transcription activator-like effectors identified in representative bacterial strains activate OsRBP11 expression. The OsRBP11 protein, in turn, facilitates alternative splicing of the OsNPR3 mRNA precursor, leading to the production of truncated OsNPR3 protein variants. The OsNPR3 variants exacerbate bacterial diseases by sequestering OsNPR1 from defense-gene activation. By contrast, both artificial and natural variations in OsRBP11 prevent the alternative splicing of OsNPR3, restore the defense function of OsNPR1, and enhance rice resistance to different bacterial strains. These findings not only reveal a novel regulatory pathway exploited by bacterial pathogens to facilitate their pathogenicity and subvert plant defense but also provide a genetic basis for biotechnological strategies aimed at developing broad-spectrum resistance in crops.

The rapid production of hydrogen peroxide (H2O2) is a hallmark of plants’ successful recognition of pathogen infection and plays a crucial role in innate immune signaling. Aquaporins (AQPs) are membrane channels that facilitate the transport of small molecular compounds across cell membranes. In plants, AQPs from the plasma membrane intrinsic protein (PIP) family are utilized for the transport of H2O2, thereby regulating various biological processes. Plants contain two PIP families, PIP1s and PIP2s. However, the specific functions and relationships between these subfamilies in plant growth and immunity remain largely unknown. In this study, we explore the synergistic role of AtPIP1;4 and AtPIP2;4 in regulating plant growth and disease resistance in Arabidopsis. We found that in plant cells treated with H2O2, AtPIP1;4 and AtPIP2;4 act as facilitators of H2O2 across membranes and the translocation of externally applied H2O2 from the apoplast to the cytoplasm. Moreover, AtPIP1;4 and AtPIP2;4 collaborate to transport bacterial pathogens and flg22-induced apoplastic H2O2 into the cytoplasm, leading to increased callose deposition and enhanced defense gene expression to strengthen immunity. These findings suggest that AtPIP1;4 and AtPIP2;4 cooperatively mediate H2O2 transport to regulate plant growth and immunity.

Type-3 effectors (T3E) of phytopathogenic Gram-negative bacteria fulfill a virulent role, causing disease, or an avirulent role, inducing immunity, following their translocation into plant cells. This study aimed to validate the hypothesis that bacterial T3E translocation requires lipidic compounds in plant cell membranes. Based on genetic, molecular, and biochemical assays, we determined that phosphatidylinositol 4-phosphate (PI4P) associated with plant cell membranes is essential for the translocation of T3E by bacterial pathogens. Replicate experimental data revealed that PI4P cooperates with the type-3 translocase HrpF to facilitate the translocation of effectors TAL and Xop from Xanthomonas oryzae and Hop from Pseudomonas syringae into the cells of Oryza sativa and Nicotiana benthamiana, respectively. Genetic and molecular analyses confirmed that, once translocated into plant cells, the distinct effectors induce disease or immunity. Combined genetic and pharmacological analyses revealed that when PI4P content is suppressed via genetic or pharmacological measures, the T3 effector translocation is considerably suppressed, resulting in serious inhibition of bacterial infection. Overall, these findings demonstrate that cooperative functioning of HrpF-PI4P is conserved in bacterial effectors and plants.

The plant signaling pathway that regulates pathogen-associated molecular pattern (PAMP)-triggered immunity (PTI) involves mitogen-activated protein kinase (MAPK) cascades that comprise sequential activation of several protein kinases and the ensuing phosphorylation of MAPKs, which activate transcription factors (TFs) to promote downstream defense responses. To identify plant TFs that regulate MAPKs, we investigated TF-defective mutants of Arabidopsis thaliana and identified MYB44 as an essential constituent of the PTI pathway. MYB44 confers resistance against the bacterial pathogen Pseudomonas syringae by cooperating with MPK3 and MPK6. Under PAMP treatment, MYB44 binds to the promoters of MPK3 and MPK6 to activate their expression, leading to phosphorylation of MPK3 and MPK6 proteins. In turn, phosphorylated MPK3 and MPK6 phosphorylate MYB44 in a functionally redundant manner, thus enabling MYB44 to activate MPK3 and MPK6 expression and further activate downstream defense responses. Activation of defense responses has also been attributed to activation of EIN2 transcription by MYB44, which has previously been shown to affect PAMP recognition and PTI development. AtMYB44 thus functions as an integral component of the PTI pathway by connecting transcriptional and posttranscriptional regulation of the MPK3/6 cascade.

Ethylene Insensitive 2 (EIN2) is an integral membrane protein that regulates ethylene signaling towards plant development and immunity by release of its carboxy-terminal functional portion (EIN2C) into the nucleus. The present study elucidates that the nuclear trafficking of EIN2C is induced by importin β1, which triggers the phloem-based defense (PBD) against aphid infestations in Arabidopsis. In plants, IMPβ1 interacts with EIN2C to facilitate EIN2C trafficking into the nucleus, either by ethylene treatment or by green peach aphid infestation, to confer EIN2-dependent PBD responses, which, in turn, impede the phloem-feeding activity and massive infestation by the aphid. In Arabidopsis, moreover, constitutively expressed EIN2C can complement the impβ1 mutant regarding EIN2C localization to the plant nucleus and the subsequent PBD development in the concomitant presence of IMPβ1 and ethylene. As a result, the phloem-feeding activity and massive infestation by green peach aphid were highly inhibited, indicating the potential value of EIN2C in protecting plants from insect attacks.

Transcription activator-like effectors (TALEs) produced by plant pathogenic bacteria mainly belonging to the genus Xanthomonas cause plant diseases through activation of host susceptibility genes in plant cell nuclei. How TALEs enter plant cell nuclei was not clear until recent studies about PthXo1 and TALI, two TALEs produced by Xanthomonas oryzae pv. oryzae ( Xoo ) and X. oryzae pv. oryzicola ( Xoc ), the rice ( Oryza sativa ) pathogens that cause bacterial blight and bacterial leaf streak, respectively. Here, we report that rice importin IMPα1b serves as a nuclear transport receptor in rice plants to facilitate the nuclear import of PthXo1 and TALI from Xoo and Xoc , respectively. While wild-type (WT) rice plants support the nuclear import of PthXo1 and TALI, nuclear trafficking is defective in OsIMPα1b loss-of-function mutants generated by clustered regularly interspaced short palindromic repeats (CRISPR)–CRISPR-associated 9 (Cas9)-based gene editing. In the edited plants infected by Xoo , OsIMPα1b sequesters PthXo1 from the nucleus, the PthXo1-targeted rice susceptibility gene is no longer activated, and bacterial virulence and blight disease are alleviated as a result. In the edited plants infected by Xoc , OsIMPα1b sequesters TALI from the nucleus, the role of TALI in suppressing rice defense responses is nullified, and rice defense responses are in turn activated to inhibit bacterial virulence and alleviate bacterial leaf streak severity.

Vesicular trafficking is a conserved material transport process in eukaryotic cells. The GGA family proteins are clathrin adaptors that are involved in eukaryotic vesicle transport, but their functions in phytopathogenic filamentous fungi remain unexplored. Here, we examined the only GGA family protein in Fusarium graminearum, FgGga1, which localizes to both the late Golgi and endosomes. In the absence of FgGga1, the fungal mutant exhibited defects in vegetative growth, DON biosynthesis, ascospore discharge and virulence. Fluorescence microscopy analysis revealed that FgGga1 is associated with trans-Golgi network (TGN)-to-plasma membrane, endosome-to-TGN and endosome-to-vacuole transport. Mutational analysis on the five domains of FgGga1 showed that the VHS domain was required for endosome-to-TGN transport while the GAT167-248 and the hinge domains were required for both endosome-to-TGN and endosome-to-vacuole transport. Importantly, the deletion of the FgGga1 domains that are required in vesicular trafficking also inhibited vegetative growth and virulence of F. graminearum. In addition, FgGga1 interacted with the ascospore discharge regulator Ca2+ ATPase FgNeo1, whose transport to the vacuole is dependent on FgGga1-mediated endosome-to-vacuole transport. Our results suggest that FgGga1 is required for fungal development and virulence via FgGga1-mediated vesicular trafficking, and FgGga1-mediated endosome-to-vacuole transport facilitates ascospore discharge in F. graminearum.

Eukaryotic aquaporins share the characteristic of functional multiplicity in transporting distinct sub-strates and regulating various processes, but the underlying molecular basis for this is largely unknown. Here, we report that the wheat (Triticum aestivum) aquaporin TaPIP2;10 undergoes phosphorylation to promote photosynthesis and productivity and to confer innate immunity against pathogens and a gener-alist aphid pest. In response to elevated atmospheric CO2 concentrations, TaPIP2;10 is phosphorylated at the serine residue S280 and thereafter transports CO2 into wheat cells, resulting in enhanced photosyn-thesis and increased grain yield. In response to apoplastic H2O2 induced by pathogen or insect attacks, TaPIP2;10 is phosphorylated at S121 and this phosphorylated form transports H2O2 into the cytoplasm, where H2O2 intensifies host defenses, restricting further attacks. Wheat resistance and grain yield could be simultaneously increased by TaPIP2;10 overexpression or by expressing a TaPIP2;10 phosphomimic with aspartic acid substitutions at S121 and S280, thereby improving both crop productivity and immunity.

To overcome pathogen infection, plants deploy a highly efficient innate immune system, which often uses hydrogen peroxide (H2O2), a versatile reactive oxygen species, to activate downstream defense responses. H2O2 is a potential substrate of aquaporins (AQPs), the membrane channels that facilitate the transport of small compounds across plasma membranes or organelle membranes. To date, however, the functional relationship between AQPs and H2O2 in plant immunity is largely undissected. Here, we report that the rice (Oryza sativa) AQP OsPIP2;2 transports pathogen-induced apoplastic H2O2 into the cytoplasm to intensify rice resistance against various pathogens. OsPIP2;2-transported H2O2 is required for microbial molecular pattern flg22 to activate the MAPK cascade and to induce the downstream defense responses. In response to flg22, OsPIP2;2 is phosphorylated at the serine residue S125, and therefore gains the ability to transport H2O2. Phosphorylated OsPIP2;2 also triggers the translocation of OsmaMYB, a membrane-anchored MYB transcription factor, into the plant cell nucleus to impart flg22-induced defense responses against pathogen infection. On the contrary, if OsPIP2;2 is not phosphorylated, OsmaMYB remains associated with the plasma membrane, and plant defense responses are no longer induced. These results suggest that OsPIP2;2 positively regulates plant innate immunity by mediating H2O2 transport into the plant cell and mediating the translocation of OsmaMYB from plasma membrane to nucleus.

Fusarium graminearum is a plant filamentous pathogenic fungi and the predominant causal agent of Fusarium head blight (FHB) in cereals worldwide. The regulators of the secretory pathway contribute significantly to fungal mycotoxin synthesis, development, and virulence. However, their roles in these processes in F. graminearum remain poorly understood. Here, we identified and functionally characterized the endoplasmic reticulum (ER) cargo receptor FgErv14 in F. graminearum. Firstly, it was observed that FgErv14 is mainly localized in the ER. Then, we constructed the FgErv14 deletion mutant (ΔFgerv14) and found that the absence of the FgErv14 caused a serious reduction in vegetative growth, significant defects in asexual and sexual reproduction, and severely impaired virulence. Furthermore, we found that the ΔFgerv14 mutant exhibited a reduced expression of TRI genes and defective toxisome generation, both of which are critical for deoxynivalenol (DON) biosynthesis. Importantly, we found the green fluorescent protein (GFP)-tagged FgRud3 was dispersed in the cytoplasm, whereas GFP-FgSnc1-PEM was partially trapped in the late Golgi in ΔFgerv14 mutant. These results demonstrate that FgErv14 mediates anterograde ER-to-Golgi transport as well as late secretory Golgi-to-Plasma membrane transport and is necessary for DON biosynthesis, asexual and sexual reproduction, vegetative growth, and pathogenicity in F. graminearum.

Plant aquaporins are a recently noted biological resource with a great potential to improve crop growth and defense traits. Here, we report the functional modulation of the rice (Oryza sativa) aquaporin OsPIP1;3 to enhance rice photosynthesis and grain production and to control bacterial blight and leaf streak, the most devastating worldwide bacterial diseases in the crop. We characterize OsPIP1;3 as a physiologically relevant CO2-transporting facilitator, which supports 30% of rice photosynthesis on average. This role is nullified by interaction of OsPIP1;3 with the bacterial protein Hpa1, an essential component of the Type III translocon that supports translocation of the bacterial Type III effectors PthXo1 and TALi into rice cells to induce leaf blight and streak, respectively. Hpa1 binding shifts OsPIP1;3 from CO2 transport to effector translocation, aggravates bacterial virulence, and blocks rice photosynthesis. On the contrary, the external application of isolated Hpa1 to rice plants effectively prevents OsPIP1;3 from interaction with Hpa1 secreted by the bacteria that are infecting the plants. Blockage of the OsPIP1;3-Hpa1 interaction reverts OsPIP1;3 from effector translocation to CO2 transport, abrogates bacterial virulence, and meanwhile induces defense responses in rice. These beneficial effects can combine to enhance photosynthesis by 29-30%, reduce bacterial disease by 58-75%, and increase grain yield by 11-34% in different rice varieties investigated in small-scale field trials conducted during the past years. Our results suggest that crop productivity and immunity can be coordinated by modulating the physiological and pathological functions of a single aquaporin to break the growth-defense tradeoff barrier.

Xanthomonas oryzae pv. oryzicola (Xoc) causes bacterial leaf streak in rice. To promote disease development, bacteria deliver effectors such as transcription activator-like (TAL) effectors into plant cells to modulate host defense and promote bacterial growth. In this study, we constructed a Tn5 transposon-mediated mutation library of GD41, which possesses 30 TAL effectors. A TalI gene deficient mutant GD21 was obtained. GD21 demonstrated compromised bacterial virulence and growth. Sequence analysis showed that TalI is a typical TAL effector gene with 14.5 repeats and RVDs identical to Tal3 in Xoc BXOR1. Expression of TalI in Xo strains resulted in enhanced bacterial growth in rice and medium. Target gene screening results suggest that TalI does not target the rice gene OsSULTR3;6 and Os09g29100. Orthology analysis showed that TalI does not belong to the 13 conserved TAL ortholog families crossing Xoc strains. Next, the role of TalI in SA, JA, and ET hormone pathways was tested, and our results showed that TalI has an effect on the expression of PR1a, PR1b, and PR5 but not on other marker genes. The HR suppression ability of TalI was then evaluated by introducing avrXa10 and avrXa7 into GD41 and GD21. The transformed strains displayed similar HR eliciting pattern on rice lines IRBB10 and IRBB7, which suggests that TalI does not interfere with effector mediated defense directly.

Plants employ aquaporins (AQPs) of the plasma membrane intrinsic protein (PIP) family to import environmental substrates, thereby affecting various processes, such as the cellular responses regulated by the signaling molecule hydrogen peroxide (H2O2). Common wheat (Triticum aestivum) contains 24 candidate members of the PIP family, designated as TaPIP1;1 to TaPIP1;12 and TaPIP2;1 to TaPIP2;12. None of these TaPIP candidates have been characterized for substrate selectivity or defense responses in their source plant. Here, we report that T. aestivum AQP TaPIP2;10 facilitates the cellular uptake of H2O2 to confer resistance against powdery mildew and Fusarium head blight, two devastating fungal diseases in wheat throughout the world. In wheat, the apoplastic H2O2 signal is induced by fungal attack, while TaPIP2;10 is stimulated to translocate this H2O2 into the cytoplasm, where it activates defense responses to restrict further attack. TaPIP2;10-mediated transport of H2O2 is essential for pathogen-associated molecular pattern-triggered plant immunity (PTI). Typical PTI responses are induced by the fungal infection and intensified by overexpression of the TaPIP2;10 gene. TaPIP2;10 overexpression causes a 70% enhancement in wheat resistance to powdery mildew and an 86% enhancement in resistance to Fusarium head blight. By reducing the disease severities, TaPIP2;10 overexpression brings about >37% increase in wheat grain yield. These results verify the feasibility of using an immunity-relevant AQP to concomitantly improve crop productivity and immunity.

In rice (Oryza sativa), the PLASMA MEMBRANE INTRINSIC PROTEIN (PIP) family of aquaporin has 11 members, OsPIP1;1 to OsPIP1;3, and OsPIP2;1 to OsPIP2;8, which are hypothesized to facilitate the transport of H2O and other small compounds across cell membranes. To date, however, only OsPIP1;2, OsPIP2;1, and OsPIP2;4 have been demonstrated for substrate selectivity in their source plant (rice). In this study, OsPIP2;2 was characterized as the most efficient facilitator of H2O transport across cell membranes in comparison with the other 10 OsPIPs. In concomitant tests of all OsPIPs, four genes (OsPIP1;3, OsPIP2;1, OsPIP2;2, and OsPIP2;4) were induced to express in leaves of rice plants following a physiological drought stress, while OsPIP2;2 was expressed to the highest level. After de novo expression in frog oocytes and yeast cells, the four OsPIP proteins were localized to the plasma membranes in trimer and tetramer and displayed the activity to increase the membrane permeability to H2O. In comparison, OsPIP2;2 was most supportive to H2O import to oocytes and yeast cells. After de novo expression in tobacco protoplasts, OsPIP2;2 exceeded OsPIP1;3, OsPIP2;1, and OsPIP2;4 to support H2O transport across the plasma membranes. OsPIP2;2-mediated H2O transport was accompanied by drought-tolerant responses, including increases in concentrations of proline and polyamines, both of which are physiological markers of drought tolerance. In rice protoplasts, H2O transport and drought-tolerant responses, which included expression of marker genes of drought tolerance pathway, were considerably enhanced by OsPIP2;2 overexpression but strongly inhibited by the gene silencing. Furthermore, OsPIP2;2 played a role in maintenance of the cell membrane integrity and effectively protected rice cells from electrolyte leakage caused by the physiological drought stress. These results suggest that OsPIP2;2 is a predominant facilitator of H2O transport in relevance to drought tolerance in the plant.

Many species of plant-pathogenic gram-negative bacteria deploy the type III (T3) secretion system to secrete virulence components, which are mostly characteristic of protein effectors targeting the cytosol of the plant cell following secretion. Xanthomonas oryzae pv. oryzae (Xoo), a rice pathogen causing bacterial blight disease, uses the T3 accessory protein HrpE to assemble the pilus pathway, which in turn secretes transcription activator-like (TAL) effectors. The hrpE gene can execute extensive physiological and pathological functions beyond effector secretion. As evidenced in this study, when the hrpE gene was deleted from the Xoo genome, the bacteria incur seriouimpairments in multiplication, motility, and virulence. The virulence nullification is attributed to reduced secretion and translocation of PthXo1, which is a TAL effector that determines the bacterial virulence in the susceptible rice varieties. When the HrpE protein produced by prokaryotic expression is applied to plants, the recombinant protein is highly effective at inducing the defense response. Moreover, leaf photosynthesis efficiency is enhanced in HrpE-treated plants. These results provide experimental avenues to modulate the plant defense and growth tradeoff by manipulating a bacterial T3 accessory protein.