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Green leaf volatiles (GLVs) play pivotal roles in plant anti-herbivore defense. This study investigated whether the rice 13-lipoxygense gene OsRCI-1 is involved in GLV production and plant defense in rice. The overexpression of OsRCI-1 (oeRCI lines) in rice resulted in increased wound-induced levels of two prominent GLVs, cis-3-hexen-1-ol and cis-3-hexenal. In a previous study, we found that the overexpression of OsRCI-1 reduced the colonization by the rice brown planthopper (BPH, Nilaparvata lugens) but increased the attractiveness to the egg parasitoid Anagrus nilaparvatae compared to wild-type (WT) plants. This study found that when cis-3-hexen-1-ol, but not cis-3-hexenal, was added to WT plants, it could change the BPH's colonization preference, i.e., more BPHs preferred to colonize the oeRCI lines. The exogenous application of cis-3-hexen-1-ol or cis-3-hexenal to BPH-infested WT plants could weaken or overturn the preference of A. nilaparvatae for oeRCI lines. However, field experiments revealed that only cis-3-hexenal was attractive to the parasitoid and increased the parasitism rates of BPH eggs. These results indicate that OsRCI-1 is involved in rice GLV production and therefore modulates both direct and indirect defense in rice. [ABSTRACT FROM AUTHOR]

Green leaf volatiles (GLVs) play pivotal roles in plant anti-herbivore defense. This study investigated whether the rice 13-lipoxygense gene OsRCI-1 is involved in GLV production and plant defense in rice. The overexpression of OsRCI-1 (oeRCI lines) in rice resulted in increased wound-induced levels of two prominent GLVs, cis-3-hexen-1-ol and cis-3-hexenal. In a previous study, we found that the overexpression of OsRCI-1 reduced the colonization by the rice brown planthopper (BPH, Nilaparvata lugens) but increased the attractiveness to the egg parasitoid Anagrus nilaparvatae compared to wild-type (WT) plants. This study found that when cis-3-hexen-1-ol, but not cis-3-hexenal, was added to WT plants, it could change the BPH’s colonization preference, i.e., more BPHs preferred to colonize the oeRCI lines. The exogenous application of cis-3-hexen-1-ol or cis-3-hexenal to BPH-infested WT plants could weaken or overturn the preference of A. nilaparvatae for oeRCI lines. However, field experiments revealed that only cis-3-hexenal was attractive to the parasitoid and increased the parasitism rates of BPH eggs. These results indicate that OsRCI-1 is involved in rice GLV production and therefore modulates both direct and indirect defense in rice.

Green leaf volatiles (GLVs) are organic compounds emitted by plants in response to insect attacks, offering early detection potential. Current GLV detection methods like gas chromatography-mass spectroscopy (GC-MS) are costly and complex and lack real-time monitoring capability. There is an unmet need for affordable and portable sensors with high sensitivity to monitor GLVs in real time. In this study, we developed a novel sensor capable of capturing piezoelectric and colorimetric signals for the sensitive and selective detection of 1-hexanol, a well-known green leaf volatile. We used a piezoelectric micro quartz tuning fork (MQTF) as the multifunctional transducer. The MQTF's two prongs were coated with a metal-organic framework (MOF)-thymol blue hybrid sensing material, enabling detection through both color change and resonating frequency shift upon 1-hexanol binding. MOFs offer a high surface area and tunable pore size, which enhance sensor sensitivity and selectivity. The sensor's frequency shift indicates mass change due to 1-hexanol binding to MOFs, while the colorimetric sensing signal relies on thymol blue's reaction with 1-hexanol. Our test results demonstrate the sensor's ability to detect 1-hexanol from 62.5 ppb to 250 ppm with high sensitivity and selectivity when the colorimetric and piezoelectric sensing signals are integrated. Due to its compact size, affordability, easy fabrication, wide detection range, and high sensitivity and selectivity, this colorimetric-piezoelectric sensor could serve as an effective tool for early detection of insect herbivore attacks and timely crop protection strategies.

Green leaf volatiles (GLVs) cover a group of mainly C6-and C9-aldehydes, -alcohols and -esters. Their name refers to their characteristic herbal and fruity scent, which is similar to that of freshly cut grass or vegetables. Lipoxygenases (LOXs) catalyze the peroxidation of unsaturated fatty acids. The resulting hydroperoxy fatty acids are then cleaved into aldehydes and oxo acids by fatty acid hydroperoxide lyases (HPLs). Herein, we equipped the yeast Komagataella phaffii with recombinant genes coding for LOX and HPL, to serve as a biocatalyst for GLV production. We expressed the well-known 13S-specific LOX gene from Pleurotus sapidus and a compatible HPL gene from Medicago truncatula. In bioconversions, glycerol induced strains formed 12.9 mM hexanal using whole cells, and 8 mM hexanol was produced with whole cells induced by methanol. We applied various inducible and constitutive promoters in bidirectional systems to influence the final ratio of LOX and HPL proteins. By implementing these recombinant enzymes in Komagataella phaffii, challenges such as biocatalyst supply and lack of product specificity can finally be overcome.

Green leaf volatiles (GLVs), the common constituents of herbivore-infested plant volatiles (HIPVs), play an important role in plant defense and function as chemical cues to communicate with other individuals in nature. Reportedly, in addition to endogenous GLVs, the absorbance of airborne GLVs emitted by infested neighboring plants also play a major role in plant defense. For example, the exclusive accumulation of (Z)-3-hexenyl vicianoside in the HIPV-exposed tomato plants occurs by the glycosylation of airborne (Z)-3-hexenol (Z3HOL); however, it is unclear how plants process the other absorbed GLVs. This study demonstrates that tomato plants dominantly accumulated GLV–glycosides after exposure to green leaf alcohols [Z3HOL, (E)-2-hexenol, and n -hexanol] using non-targeted LC–MS analysis. Three types of green leaf alcohols were independently glycosylated without isomerization or saturation/desaturation. Airborne green leaf aldehydes and esters were also glycosylated, probably through converting aldehydes and esters into alcohols. Further, we validated these findings in Arabidopsis mutants- (Z)-3-hexenal (Z3HAL) reductase (chr) mutant that inhibits the conversion of Z3HAL to Z3HOL and the acetyl-CoA:(Z)-3-hexen-1-ol acetyltransferase (chat) mutant that impairs the conversion of Z3HOL to (Z)-3-hexenyl acetate. Exposure of the chr and chat mutants to Z3HAL accumulated lower and higher amounts of glycosides than their corresponding wild types (Col-0 and L er), respectively. These findings suggest that plants process the exogenous GLVs by the reductase(s) and the esterase(s), and a part of the processed GLVs contribute to glycoside accumulation. Overall, the study provides insights into the understanding of the communication of the plants within their ecosystem, which could help develop strategies to protect the crops and maintain a balanced ecosystem. [ABSTRACT FROM AUTHOR]

Green leaf volatiles (GLVs), the common constituents of herbivore-infested plant volatiles (HIPVs), play an important role in plant defense and function as chemical cues to communicate with other individuals in nature. Reportedly, in addition to endogenous GLVs, the absorbance of airborne GLVs emitted by infested neighboring plants also play a major role in plant defense. For example, the exclusive accumulation of (Z)-3-hexenyl vicianoside in the HIPV-exposed tomato plants occurs by the glycosylation of airborne (Z)-3-hexenol (Z3HOL); however, it is unclear how plants process the other absorbed GLVs. This study demonstrates that tomato plants dominantly accumulated GLV–glycosides after exposure to green leaf alcohols [Z3HOL, (E)-2-hexenol, and n-hexanol] using non-targeted LC–MS analysis. Three types of green leaf alcohols were independently glycosylated without isomerization or saturation/desaturation. Airborne green leaf aldehydes and esters were also glycosylated, probably through converting aldehydes and esters into alcohols. Further, we validated these findings in Arabidopsis mutants- (Z)-3-hexenal (Z3HAL) reductase (chr) mutant that inhibits the conversion of Z3HAL to Z3HOL and the acetyl-CoA:(Z)-3-hexen-1-ol acetyltransferase (chat) mutant that impairs the conversion of Z3HOL to (Z)-3-hexenyl acetate. Exposure of the chr and chat mutants to Z3HAL accumulated lower and higher amounts of glycosides than their corresponding wild types (Col-0 and Ler), respectively. These findings suggest that plants process the exogenous GLVs by the reductase(s) and the esterase(s), and a part of the processed GLVs contribute to glycoside accumulation. Overall, the study provides insights into the understanding of the communication of the plants within their ecosystem, which could help develop strategies to protect the crops and maintain a balanced ecosystem.

Herbivore-induced plant volatiles (HIPVs) provide direct benefits to plants as antimicrobials and herbivore repellents, but their potential as direct toxins to herbivores is unclear. Here, we assayed the larvicidal activity of six common HIPVs from three different biochemical pathways and tested the hypothesis that the larvicidal activity of HIPVs is related to the host specialization of the insect pest. We first assessed β-caryophyllene, linalool, z-3-hexenyl acetate, z-3-hexenol, e-2-hexenal, and indole against the beet armyworm (Spodoptera exigua) and found that indole was sevenfold more toxic compared to the other volatiles when incorporated into the diet. Then, we tested the larvicidal activity of indole against six common, destructive pest caterpillars with varying host ranges. Consistent with our hypothesis, indole toxicity varied with caterpillar host range: indole toxicity was sevenfold higher in more specialized insect species relative to generalist insect species. That said, the LC50 of indole was comparable to other reported anti-herbivore agents even against the generalist caterpillars. Yet, indole in headspace had neither larvicidal nor ovicidal activity on any caterpillar species tested. These results support a key ecological precept of a trade-off between host specialization and chemical detoxification and also indicate that indole in particular is directly toxic to herbivores and therefore potentially useful in integrated pest management strategies. [ABSTRACT FROM AUTHOR]

The timing of plant volatile emissions is important for a robust indirect defense response. Green leaf volatiles (GLVs) are emitted by plants upon damage but can be suppressed by herbivore‐associated elicitors, and the abundance and composition of GLVs vary depending on the timing of herbivore attack. We show that the GLV biosynthetic enzyme HYDROPEROXIDE LYASE (HPL) is transcriptionally regulated by the circadian clock in Nicotiana attenuata. In accordance with transcript abundance of NaHPL, GLV aldehyde pools in intact leaves peaked at night and at subjective night under diurnal and continuous light conditions, respectively. Moreover, although the basal abundance of NaHPL transcripts is upregulated by jasmonate (JA) signaling, JA does not regulate the reduction of NaHPL transcript abundance in damaged leaves by simulated herbivore treatment. Unexpectedly, the plant circadian clock was strongly altered when Manduca sexta larvae fed on N. attenuata, and this was also independent of JA signaling. Lastly, the temporal dynamics of NaHPL transcripts and total GLV emissions were strongly altered by M. sexta larval feeding. Our data suggest that the temporal dynamics of emitted GLV blends result from a combination of damage, JA signaling, herbivore‐associated elicitors, and the plant circadian clock. Herbivore‐induced plant volatiles display complex temporal dynamics, which matter for effectively attracting the natural enemies of herbivores. Here, we show that green leaf volatile (GLV) biosynthesis in the wild tobacco Nicotiana attenuata has a circadian rhythm. Interestingly, although the biosynthesis of GLVs is jasmonate‐dependent, herbivore elicitors suppress GLV biosynthesis. Moreover, herbivory strongly alters the diurnal rhythm of circadian clock gene transcript levels: the effect of herbivory differs depending on the elicitors used and treatment time. This study reveals how a plant shapes its temporal dynamics of GLVs emissions by integrating circadian clock signaling with herbivore‐induced responses. [ABSTRACT FROM AUTHOR]

Summary: Plants emit a plethora of volatile organic compounds in response to biotic and abiotic stresses. These compounds act as infochemicals for ecological communication in the phytobiome. This study reviews the role of microbe‐induced plant volatiles (MIPVs) in plant–microbe interactions. MIPVs are affected by the taxonomic position of the microbe, the identity of the plant and the type of interaction. Plants also emit exclusive blends of volatiles in response to nonhost and host interactions, as well as to beneficial microbes and necrotrophic/biotrophic pathogens. These MIPVs directly inhibit pathogen growth and indirectly promote resistance/susceptibility to subsequent plant pathogen attack. Viruses and phloem‐limiting bacteria modify plant volatiles to attract insect vectors. Susceptible plants can respond to MIPVs from resistant plants and become resistant. Recent advances in our understanding of the molecular mechanisms of MIPV synthesis in plants and how plant pathogen effectors manipulate their biosynthesis are discussed. This knowledge will help broaden our understanding of plant–microbe interactions and should facilitate the development of new emerging techniques for sustainable plant disease management. See also the Editorial by Kessler, 220: 655–658. [ABSTRACT FROM AUTHOR]

Synechocystis salt-responsive gene 1 (sysr1) was engineered for expression in higher plants, and gene construction was stably incorporated into tobacco plants. We investigated the role of Sysr1 [a member of the alcohol dehydrogenase (ADH) superfamily] by examining the salt tolerance of sysr1-overexpressing (sysr1-OX) tobacco plants using quantitative real-time polymerase chain reactions, gas chromatography-mass spectrometry, and bioassays. The sysr1-OX plants exhibited considerably increased ADH activity and tolerance to salt stress conditions. Additionally, the expression levels of several stress-responsive genes were upregulated. Moreover, airborne signals from salt-stressed sysr1-OX plants triggered salinity tolerance in neighboring wild-type (WT) plants. Therefore, Sysr1 enhanced the interconversion of aldehydes to alcohols, and this occurrence might affect the quality of green leaf volatiles (GLVs) in sysr1-OX plants. Actually, the Z-3-hexenol level was approximately twofold higher in sysr1-OX plants than in WT plants within 1–2 h of wounding. Furthermore, analyses of WT plants treated with vaporized GLVs indicated that Z-3-hexenol was a stronger inducer of stress-related gene expression and salt tolerance than E-2-hexenal. The results of the study suggested that increased C6 alcohol (Z-3-hexenol) induced the expression of resistance genes, thereby enhancing salt tolerance of transgenic plants. Our results revealed a role for ADH in salinity stress responses, and the results provided a genetic engineering strategy that could improve the salt tolerance of crops.

Green leaf volatiles (GLVs) have been reported to play an important role in the host-locating behavior of several folivores that feed on angiosperms. However, next to nothing is known about how the green leafhopper, Empoasca vitis, chooses suitable host plants and whether it detects differing emission levels of GLV components among genetically different tea varieties. Here we found that the constitutive transcript level of the tea hydroperoxide lyase (HPL) gene CsiHPL1, and the amounts of (Z)-3-hexenyl acetate and of total GLV components are significantly higher in tea varieties that are susceptible to E. vitis (Enbiao (EB) and Banzhuyuan (BZY)) than in varieties that are resistant to E. vitis (Changxingzisun (CX) and Juyan (JY)). Moreover, the results of a Y-tube olfactometer bioassay and an oviposition preference assay suggest that (Z)-3-hexenyl acetate and (Z)-3-hexenol offer host and oviposition cues for E. vitis female adults. Taken together, the two GLV components, (Z)-3-hexenol and especially (Z)-3-hexenyl acetate, provide a plausible mechanism by which tea green leafhoppers distinguish among resistant and susceptible varieties. Future research should be carried out to obtain the threshold of the above indices and then assess their reasonableness. The development of practical detection indices would greatly improve our ability to screen and develop tea varieties that are resistant to E. vitis. [ABSTRACT FROM PUBLISHER]

Background: The dynamics of plant volatile (PV) emission, and the relationship between damaged area and biosynthesis of bioactive molecules in plant-insect interactions, remain open questions. Direct Contact-Sorptive Tape Extraction (DC-STE) is a sorption sampling technique employing non adhesive polydimethylsiloxane tapes, which are placed in direct contact with a biologically-active surface. DC-STE coupled to Gas Chromatography-Mass Spectrometry (GC-MS) is a non-destructive, high concentration-capacity sampling technique able to detect and allow identification of PVs involved in plant responses to biotic and abiotic stresses. Here we investigated the leaf topographical dynamics of herbivory-induced PV (HIPV) produced by Phaseolus lunatus L. (lima bean) in response to herbivory by larvae of the Mediterranean climbing cutworm (Spodoptera littoralis Boisd.) and mechanical wounding by DC-STE-GC-MS. Results: Time-course experiments on herbivory wounding caused by larvae (HW), mechanical damage by a pattern wheel (MD), and MD combined with the larvae oral secretions (OS) showed that green leaf volatiles (GLVs) [(E)-2-hexenal, (Z)-3-hexen-1-ol, 1-octen-3-ol, (Z)-3-hexenyl acetate, (Z)-3-hexenyl butyrate] were associated with both MD and HW, whereas monoterpenoids [(E)-β-ocimene], sesquiterpenoids [(E)-nerolidol] and homoterpenes (DMNT and TMTT) were specifically associated with HW. Up-regulation of genes coding for HIPV-related enzymes (Farnesyl Pyrophosphate Synthase, Lipoxygenase, Ocimene Synthase and Terpene Synthase 2) was consistent with HIPV results. GLVs and sesquiterpenoids were produced locally and found to influence their own gene expression in distant tissues, whereas (E)-β-ocimene, TMTT, and DMNT gene expression was limited to wounded areas. Conclusions: DC-STE-GC-MS was found to be a reliable method for the topographical evaluation of plant responses to biotic and abiotic stresses, by revealing the differential distribution of different classes of HIPVs. The main advantages of this technique include: a) in vivo sampling; b) reproducible sampling; c) ease of execution; d) simultaneous assays of different leaf portions, and e) preservation of plant material for further "omic" studies. DC-STE-GC-MS is also a low-impact innovative method for in situ PV detection that finds potential applications in sustainable crop management. [ABSTRACT FROM AUTHOR]

Green leaf volatiles (GLVs) cover a group of mainly C6-and C9-aldehydes, -alcohols and -esters. Their name refers to their characteristic herbal and fruity scent, which is similar to that of freshly cut grass or vegetables. Lipoxygenases (LOXs) catalyze the peroxidation of unsaturated fatty acids. The resulting hydroperoxy fatty acids are then cleaved into aldehydes and oxo acids by fatty acid hydroperoxide lyases (HPLs). Herein, we equipped the yeast Komagataella phaffii with recombinant genes coding for LOX and HPL, to serve as a biocatalyst for GLV production. We expressed the well-known 13 S -specific LOX gene from Pleurotus sapidus and a compatible HPL gene from Medicago truncatula . In bioconversions, glycerol induced strains formed 12.9 mM hexanal using whole cells, and 8 mM hexanol was produced with whole cells induced by methanol. We applied various inducible and constitutive promoters in bidirectional systems to influence the final ratio of LOX and HPL proteins. By implementing these recombinant enzymes in Komagataella phaffii , challenges such as biocatalyst supply and lack of product specificity can finally be overcome., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Hashem, Hochrinner, Bürgler, Rinnofner, Pichler and Winkler.)

In response to herbivory by cabbage white butterfly (Pieris rapae) larvae, crucifer plants emit green leaf volatiles (GLVs) that attractCotesia glomerata, a carnivorous natural enemy of the larvae. To artificially increase GLV production by crucifers, we created transgenicArabidopsisthat constitutively expressed cucumber cytosolic lipoxygenase (CsLOX2). TransgenicArabidopsis(p35S::CsLOX2.6andp35S::CsLOX2.14) infested withP. rapaelarvae were more attractive toC. glomeratathan wild type (wt) and produced more jasmonic acid.p35S::CsLOX2.14had a higher incidence of parasitism of the larvae than did wt. Infested transgenic plants emitted more (Z)-3-hexenyl acetate, (E)-β-ocimene, (Z)-β-ocimene, and DMNT than infested wt plants; (Z)-3-hexenyl acetate is known to attractCotesiawasps. Transforming cruciferous crops with cytosolicCsLOX2could aid integrated pest management via tritrophic interactions. [ABSTRACT FROM PUBLISHER]

Synechocystis salt-responsive gene 1 ( sysr1 ) was engineered for expression in higher plants, and gene construction was stably incorporated into tobacco plants. We investigated the role of Sysr1 [a member of the alcohol dehydrogenase (ADH) superfamily] by examining the salt tolerance of sysr1 -overexpressing ( sysr1 -OX) tobacco plants using quantitative real-time polymerase chain reactions, gas chromatography-mass spectrometry, and bioassays. The sysr1 -OX plants exhibited considerably increased ADH activity and tolerance to salt stress conditions. Additionally, the expression levels of several stress-responsive genes were upregulated. Moreover, airborne signals from salt-stressed sysr1 -OX plants triggered salinity tolerance in neighboring wild-type (WT) plants. Therefore, Sysr1 enhanced the interconversion of aldehydes to alcohols, and this occurrence might affect the quality of green leaf volatiles (GLVs) in sysr1 -OX plants. Actually, the Z -3-hexenol level was approximately twofold higher in sysr1 -OX plants than in WT plants within 1-2 h of wounding. Furthermore, analyses of WT plants treated with vaporized GLVs indicated that Z -3-hexenol was a stronger inducer of stress-related gene expression and salt tolerance than E -2-hexenal. The results of the study suggested that increased C 6 alcohol ( Z -3-hexenol) induced the expression of resistance genes, thereby enhancing salt tolerance of transgenic plants. Our results revealed a role for ADH in salinity stress responses, and the results provided a genetic engineering strategy that could improve the salt tolerance of crops.