The effect of a mixotrophic chrysophyte on toxic and colony-forming cyanobacteria

1. In order to test the effect of Ochromonas sp. , a mixotrophic chrysophyte, on cyanobacteria, grazing experiments were performed under controlled conditions. We studied grazing on three Microcystis aeruginosa strains, varying in toxicity and morphology, as well as on one filamentous cyanobacterium, Pseudanabaena sp. Furthermore, we analysed the co-occurrence of Ochromonas and Microcystis in natural systems in relation to various environmental parameters (TP, TN, DOC, temperature, pH), using data from 460 Norwegian lakes.
2.  Ochromonas was able to feed on all four cyanobacterial strains tested, and grew quickly on all of them. The chrysophyte caused net growth reductions in all three Microcystis strains (the very toxic single-celled strain PCC 7806; the less toxic colony-forming Bear AC and the less toxic single-celled Spring CJ). The effect of Ochromonas was strongest on the Spring CJ strain. Although the effect of Ochromonas grazing on the growth of Pseudanabaena was relatively smaller, it also reduced the net growth of this cyanobacterium significantly.
3. After 4 days of incubation with Ochromonas the total amount of cyanotoxins in the three Microcystis strains was reduced by 91.1–98.7% compared with the controls.
4.  Ochromonas occurred in similar densities across all 460 Norwegian lakes. Microcystis occurred only at higher TN, TP, temperature and pH values, although its density was often several orders of magnitude higher than that of Ochromonas . Ochromonas co-occurred in 94% of the samples in which Microcystis was present.
5. From our study it is not clear whether Ochromonas could control Microcystis blooms in natural lakes. However, our study does demonstrate that Ochromonas usually occurs in lakes with Microcystis , and our small scale experiments show that Ochromonas can strongly reduce the biomass of Microcystis and its toxin content.     

Comparative demographic impacts of `info-chemicals' and exploitative competition: an empirical test using Daphnia magna

1. Crowding `info-chemicals', metabolites released into the environment that act as signals to conspecifics as well as other organisms, have often been shown to cause phenotypic plasticity in cladoceran life-history traits including morphology, reproductive strategy and sex induction. Effects on population demography and diversity, however, have not been examined directly.
2. Replicate laboratory populations of the zooplankter, Daphnia magna, were started with 250 juveniles and allowed to develop for 8 weeks in 10 L flow-through aquaria with food levels of 1 mg CL^–1. Two crowded water treatments from different clonal sources were compared with untreated water as a control.
3. The presence of crowding chemicals affected initial population structure, through reduction in parthenogenetic female body lengths, brood sizes and juvenile densities. However, the timing of population density maxima and subsequent population structure, clonal composition and diversity were similar among all treatments.
4. Clonal richness remained slightly higher in untreated populations because of increased prevalence of rare clones.
5. Exploitation competition had greater impact on population structure than the relative concentration of crowding chemicals, indicating that density dependent effects were mediated primarily by competition for common resources.
6. Crowding info-chemicals may play a greater role in community structure than in single species population structure because of taxonomic diversity in species' response that may be unrelated to the source species density.     

Responses of Daphnia pulex populations to toxic cyanobacteria

1. We studied the effects of toxic cyanobacteria, Microcystis aeruginosa , on Daphnia pulex populations. The experiment was performed at room temperature in laboratory microcosms to which we added toxic Microcystis in two pulses. Green alga, Scenedesmus obtusiusculus, was provided in two concentrations.
2. Microcystis exposure resulted in a decreased population density of Daphnia. The proportion of adolescents was higher in the Microcystis treatment than in the control, while the proportion of newborns did not differ significantly from the control. This indicates delayed maturation of Daphnia exposed to Microcystis . We found no significant impacts of cyanobacteria, food level or clonal origin on population variability after correcting for differences in population size.
3. Juveniles of the two clones studied showed different sensitivities to toxic Microcystis in the acute test. However, in the population experiment we did not find clonal differences between Microcystis and control treatments in the relative proportion of juveniles.
4. The number of ephippia produced per adult was highest in the treatments with Microcystis. This indicates that cyanobacterial toxins may be one of several factors inducing ephippia formation, acting directly or through inhibition of feeding.     

Consumption of cyanobacteria by roach (Rutilus rutilus): useful or harmful to the fish?

1. The ability of roach to use cyanobacterial food is generally believed to be one reason for the dominance of roach over perch in eutrophic European lakes. The aim of this study was to test whether cyanobacteria really are a suitable food for juvenile roach. Special attention was paid to differences between the two cyanobacteria species Aphanizomenon and Microcystis which are common in eutrophic lakes and are ingested by roach there.
2. We performed growth and behaviour experiments with juvenile roach fed with zooplankton and the different cyanobacteria. Growth rate with Aphanizomenon was lower than with Daphnia but significantly higher than without food, whereas growth rate with Microcystis was as low as without food.
3. In cultivation experiments of roach faeces, Microcystis was found not to have been digested and grew exponentially after passing through the gut whereas Aphanizomenon stayed at low biomass. Differences in growth were not related to the toxin content of cyanobacteria. Investigations of roach motility showed no differences whether fed with Aphanizomenon or Microcystis .
4. In contrast to Microcystis , Aphanizomenon can be regarded as a suitable food source for juvenile roach probably because of its better digestability. We conclude that the ability to feed on cyanobacteria is not a general competitive advantage for roach, but the outcome depends on the species composition of the cyanobacteria.     

Grazing on toxic and non-toxic Microcystis aeruginosa PCC7820 by Unio douglasiae and Corbicula fluminea

To explore the potential grazing effects of mussels on Microcystis aeruginosa, a common bloom-forming phytoplankton, Unio douglasiae and Corbicula fluminea were fed with Scenedesmus obliquus, toxic and non-toxic strains of Microcystis aeruginosa as single food and as mixtures in the laboratory. When fed with single foods, U. douglasiae has similar clearance rates on the three algae populations, while C. fluminea has significantly lower clearance rate on toxic M. aeruginosa than those on the other two algae populations. When fed with mixture foods, both the mussels show significantly higher clearance rates than on single foods. The clearance rates of U. douglasiae on the different food mixtures are not significantly different, and C. fluminea has a significantly lower clearance rate on the toxic food mixtures than that on non-toxic food mixtures. Although the relative lower clearance rates of C. fluminea on toxic food, we may still deduce that both the mussels can exert grazing pressure on phytoplankton. The deduction is supported by the composition of the excretion products. The excretion products (faeces and pseudofaeces) of both mussels contained mainly S. obliquus. In both mixed-food treatments, the ratios of S. obliquus to M. aeruginosa in the excrete products are significantly higher than those in the foods. Therefore, it can be concluded that both mussels prefer M. aeruginosa to S. obliquus, and can cause grazing pressure on M. aeruginosa.     

Somatic and population growth in selected cladoceran and rotifer species offered the cyanobacterium it Microcystis aeruginosa as food

The ability of cladocerans and rotifers to utilise the cyanobacterium Microcystis aeruginosa was tested by comparing the somatic and population growth in cultures using Chlorella and Microcystis as food types. Five species of cladocerans (Ceriodaphnia cornuta, Scapholeberis kingi, Moina macrocopa, Daphnia carinata, Simocephalus vetulus) and two species of rotifers (Brachionus calyciflorus, Hexarthra mira) were used in this study. In order to exclude the possibility of poor utilisation of Microcystis due to mechanical interference, single cells of Microcystis, (obtained by sonicating large colonies) were also offered. Experiments were done at 20 °Cs and 30 °C . In all the treatments tested, the population growth rate per day of the cladocerans ranged from -0.715 to 0.612 and that of the rotifers from -1.15 to 0.781. While C. cornuta, S. kingi and S. vetulus could utilise Microcystis, M. macrocopa and D. carinata were extremely susceptible to its toxins. The ability of the cladoceran populations to grow on Microcystis single cells was not related to the body length or gut length alone but to their ratio. The toxic effects of Microcystis were mitigated at the higher temperature. A strain of C. cornuta, collected from a Microcystis-dominated lake, had a higher growth rate on the toxic cyanobacteria suggesting that the tolerance to Microcystis could be a heritable trait. Of the two rotifer species, only H. mira survived and reproduced in some treatments of Microcystis. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genotype-dependent interactions among sympatric Microcystis strains mediated by Daphnia grazing

Natural populations of the bloom forming cyanobacterium Microcystis are typically composed of several distinct genotypes. Using Microcystis strains that differ in growth rate, microcystin production and colony formation, we conducted a laboratory experiment in the presence and absence of a grazer, the water flea Daphnia, to investigate whether interactions among strains can be predicted from functional traits, and whether the outcome of competition between strains is influenced by a grazer. Two toxic and two non‐toxic Microcystis strains, isolated from a single lake, were grown during four weeks as single strains, in all possible combinations of two strains and all together, in the presence and absence of Daphnia magna. The relative abundance of strains in the populations was assessed using denaturing gradient gel electrophoresis, and the growth rate of each strain in mixed populations was compared to its growth rate in monoculture to determine interactions between strains. The observed interactions were strain‐specific, and the relative abundances of strains in mixed populations could be partially explained by taking toxicity and colony formation into account. Importantly, some of the interactions were strongly altered by the presence of Daphnia. Daphnia induced colony formation in one strain, which then became a better competitor. Daphnia grazing also caused a higher evenness in the populations, both through a weakening of competitive interactions as well as by facilitation effects. Strong facilitation effects were due to non‐toxic strains benefiting from the protection offered by toxic strains in the presence of predation. Overall, our results emphasize the presence of strong competitive interactions between Microcystis strains in the absence of grazing, whereas indirect positive interactions are prevalent in the presence of a generalist grazer. Our results suggest that differences in functional traits and grazer‐mediated facilitation effects may enhance coexistence of Microcystis strains, including toxic and non‐toxic strains.     

Comparing grazing by Dreissena polymorpha on phytoplankton in the presence of toxic and non-toxic cyanobacteria

SUMMARY 1. The feeding behaviour of the zebra mussel ( Dreissena polymorpha ) was studied in the laboratory on different combinations of food, including a green alga ( Chlamydomonas reinhardtii ) and toxic and non-toxic strains of the cyanobacterium Microcystis aeruginosa .
2. The highest clearance rate of phytoplankton by zebra mussels was found when the mussels were feeding on a mixture of Chlamydomonas and non-toxic Microcystis , the lowest on a mixture of Chlamydomonas and toxic Microcystis .
3. The differences found in the clearance rates between food combinations can be partly explained by the production of pseudofaeces containing live phytoplankton cells. Zebra mussels expelled significantly more live phytoplankton cells in the presence of toxic Microcystis than in the presence of non-toxic Microcystis . The pseudofaeces contained predominantly live Chlamydomonas cells. Proportionately much less live Microcystis cells were encountered in the pseudofaeces.
4. Consequently, grazing of zebra mussels on a combination of Chlamydomonas and Microcystis may finally result in a dominance of Chlamydomonas over Microcystis . The presence of toxic Microcystis may even strengthen this shift.     

Grazing of two toxic Planktothrix species by Daphnia pulicaria: potential for bloom control and transfer of microcystins

The role of zooplankton in the control of cyanobacterial bloomsand the transfer of cyanotoxins to higher trophic levels areof great importance to the management of water resources. Manystudies have focused on the cyanobacterium Microcystis, butfew have examined the interactions between zooplankton and filamentouscyanobacteria. In this study, we provide experimental evidencefor the potential grazing of two toxic strains of filamentouscyanobacteria, Planktothrix rubescens and P. agardhii, by Daphniapulicaria, and for transfer of toxins in the planktonic foodchain. We determined clearance rates (CRs) by adult and juvenileD. pulicaria of the two Planktothrix strains, Scenedesmus acutusand a mixture of S. acutus cells with P. rubescens culture filtrate.Filament lengths were analyzed, and microcystin (MCY) presencein Daphnia was assessed using the Protein Phosphatase-2A (PP-2A)Inhibition Assay. The two Planktothrix strains were equallygrazed by D. pulicaria, but at lower CRs than S. acutus. Potentialanti-grazer toxins in P. rubescens filtrate did not inhibitDaphnia grazing. Small P. rubescens (<100 µm) filamentswere preferentially grazed by adult D. pulicaria, suggestingtheir limited ability to control a Planktothrix population duringa bloom. Large quantities of MCYs were found in unstarved Daphniapreviously exposed to Planktothrix, whereas quantities weresignificantly smaller in individuals starved for 24 h beforepreservation. This indicated a potential for transfer of toxinsin the food chain by Daphnia, especially immediately after ingestionof toxic cyanobacteria.     

Periphyton as alternative food source for the filter-feeding cladoceran Daphnia magna

1.  Daphnia magna , a well-studied primary consumer, is mainly known as a filter feeder. In this study, we investigated the ability of D. magna to use periphyton as an alternative food source to phytoplankton. We examined the development of laboratory populations fed with different food sources ( Desmodesmus subspicatus and/or periphyton or neither) over a period of 42 days, and observed the behaviour of the daphnids.
2.  The addition of periphyton to phytoplankton food led to an increase of daphnid population biomass. When fed with periphyton as the only food source, a small but stable D. magna population developed.
3.  The behaviour of daphnids fed with both food sources revealed a preference for feeding on D. subspicatus . Only below a concentration of D. subspicatus of approximately 0.05 mg C L⁻¹ (0.4 × 10⁷ cells L⁻¹) did D. magna use periphyton as an alternative food source.
4.  Periphyton showed distinct reactions to grazing by D. magna . The thickness of the periphyton layer was reduced from about 4 to 1 mm and we observed a change in species composition due to grazing.
5.  The ability of D. magna to graze on periphyton could serve to stabilize its population density and reinforce its competitive advantage over other cladocerans. By switching between food sources, D. magna can act as a coupler between pelagic and benthic habitats and food webs.     

Microsatellite PCR profiling of Saccharomyces cerevisiae strains during wine fermentation

AIMS: Use of microsatellite PCR to monitor populations of Saccharomyces cerevisiae strains during fermentation of grape juice. METHOD AND RESULTS: Six commercial wine strains of S. cerevisiae were screened for polymorphism at the SC8132X locus using a modified rapid PCR identification technique. The strains formed four distinct polymorphic groups that could be readily distinguished from one another. Fermentations inoculated with mixtures of three strains polymorphic at the SC8132X locus were monitored until sugar utilization was complete, and all exhibited a changing population structure throughout the fermentation. CONCLUSIONS: Rapid population quantification demonstrated that wine fermentations are dynamic and do not necessarily reflect the initial yeast population structure. One or more yeast strains were found to dominate at different stages of the fermentation. SIGNIFICANCE AND IMPACT OF THE STUDY: The population structure of S. cerevisiae during mixed culture wine fermentation is dynamic and could modify the chemical composition and flavour profile of wine.     

A spontaneous mutant of microcystin biosynthesis: genetic characterization and effect on Daphnia

Microcystis aeruginosa strain MRC is unique in its' possession of the mcyA-J gene cluster, which encodes microcystin synthetase, but its' inability to produce microcystins. M. aeruginosa strain MRD is genetically identical to MRC at numerous genomic loci examined, but produces a variety of microcystins, mainly with the amino acid tyrosine in the molecule. Zooplankton studies with Daphnia galeata and D. pulicaria , using the mutant (MRC) and its' wild type (MRD), showed for the first time that microcystins other than microcystin-LR can be responsible for the poisoning of Daphnia by Microcystis . Regardless of microcystin content, both Daphnia exhibited significantly reduced ingestion rates when fed with either strain of M. aeruginosa compared with the green alga Scenedesmus acutus . A disruption of the molting process in both Daphnia spp. was noted when these species were fed with MRC cells. Such symptoms on Daphnia have not been previously reported for cyanobacteria and may point to a bioactive compound, other than microcystin, which inhibits the hardening of protein–chitin complexes in Daphnia .     

Initial genetic diversity enhances population establishment and alters genetic structuring of a newly established Daphnia metapopulation

When newly created habitats are initially colonized by genotypes with rapid population growth rates, later arriving colonists may be prevented from establishing. Although these priority effects have been documented in multiple systems, their duration may be influenced by the diversity of the founding population. We conducted a large‐scale field manipulation to investigate how initial clonal diversity influences temporal and landscape patterns of genetic structure in a developing metapopulation. Six genotypes of obligately asexual Daphnia pulex were stocked alone (no clonal diversity) or in combination (‘high’ clonal diversity) into newly created experimental woodland ponds. We also measured the population growth rate of all clones in the laboratory when raised on higher‐quality and lower‐quality resources. Our predictions were that in the 3 years following stocking, clonally diverse populations would be more likely to persist than nonclonally diverse populations and exhibit evidence for persistent founder effects. We expected that faster growing clones would be found in more pools and comprise a greater proportion of individuals genotyped from the landscape. Genetic composition, both locally and regionally, changed significantly following stocking. Six of 27 populations exhibited evidence for persistent founder effects, and populations stocked with ‘high’ clonal diversity were more likely to exhibit these effects than nonclonally diverse populations. Performance in the laboratory was not predictive of clonal persistence or overall dominance in the field. Hence, we conclude that although laboratory estimates of fitness did not fully explain metapopulation genetic structure, initial clonal diversity did enhance D. pulex population establishment and persistence in this system.     

Colonization of Daphnia magna in a newly created pond: founder effects and secondary immigrants

In habitats recently colonized by cyclical parthenogens, founder events lead to genetic differences between populations that do not erode quickly despite ongoing dispersal. By comparing the genetic composition during initial colonization with that of the diapausing egg bank at a local scale, we here present the relative contribution of the founding clones to the build-up of genetic diversity and differentiation of a newly established cladoceran population. We monitored the population genetic structure of Daphnia magna in one newly created pond as well as the diapausing egg banks of four water bodies in the neighbouring area. Our population was founded by four individuals. After the first growing season, the largest contribution to the sexually produced resting egg bank came from only two clones. Descendants of initially rare clones and potentially also additional immigrant clones profited from outbreeding vigour and increased their frequency during the first few years after colonization. Beyond this, no further significant changes in genetic structure were observed in the egg bank. At this point, priority effects became fully operational and led to sustained population genetic differentiation from nearby ponds. Our results support that colonization dynamics strongly influence within and among population genetic variation and evolutionary potential of populations.     

Population Dynamics of Pseudomonas syringae pv. tomato Strains on Tomato Cultivars Rio Grande and Rio Grande-Pto under Field Conditions

We examined the effects of the Pto resistance locus on the population dynamics of Pseudomonas syringae pv. tomato (Pst) strains in field experiments with the nearly isogenic tomato lines Rio Grande (RG, susceptible to Pst races 0 and 1) and Rio Grande-Pto (RG-Pto, resistant to Pst race 0, susceptible to Pst race 1). Pst strain SM78-1Sm^r (race 0) grew well under field conditions and caused ample bacterial speck disease on susceptible RG plants. In contrast, strain DC3000 failed to establish large populations when inoculated onto field grown RG plants. Mean population sizes of SM78-1Sm^r were 4–5 orders of magnitude larger on RG than RG-Pto plants indicating that RG-Pto plants were highly effective in attenuating pathogen population development. Most of the sampled leaflets from RG-Pto field plots harboured small numbers of SM78-1Sm^r. However, population sizes SM78-1Sm^r as large as 10⁵–10⁶ CFU were found on a few leaflets. Bacteria isolated from these leaflets had phenotypes characteristic of Pst race 1 strains. In growth chamber plant assays, the bacterial strains grew well and caused typical speck lesions on RG-Pto plants. The strains appeared to be race-shift mutants of SM SM78-1Sm^r. Interestingly, results from DNA hybridization experiments demonstrated that the race-shift mutants were deleted for the avirulence gene, avrPto but not for avrPtoB.     

Predation and priority effects in experimental zooplankton communities

During colonization of new habitats, the sequence of arrival among species is in many cases determined by chance. Priority effects imply that differences in arrival time may lead to long-lasting differences in species dominance. To evaluate the importance of priority effects, we performed a community assembly experiment, manipulating the inoculation order of three large cladoceran zooplankton species. The inoculation treatments were crossed with a predation treatment to test whether the presence of a predator (larvae of the phantom midge Chaoborus ) results in changes in the strength of species sorting and priority effects. Our results clearly demonstrate priority effects, but also that the presence of a predator impacts both community assembly and the strength of priority effects. In the predation-free treatments, communities were dominated by either Daphnia magna or Daphnia obtusa , depending on the species arrival sequence. Whenever D. obtusa was inoculated after D. magna , the species displayed negative growth. In the presence of Chaoborus predation, the communities were generally dominated by the third species, Simocephalus vetulus . Here, the growth of D. magna was negative when the species was inoculated as second or third. Overall, our results underscore the importance of both priority effects and species sorting during community assembly.     

Inoculation history affects community composition in experimental freshwater bacterioplankton communities

Priority effects occur when the arrival order of species or genotypes has a lasting effect on community or population structure. For freshwater bacteria , priority effects have been shown experimentally among individual species, but no experiments have been performed using complex natural communities. We investigated experimentally whether a foreign bacterioplankton community influences the community assembly trajectory when inoculated prior to the local community, whether inoculation time lag affects priority effects, and how the individual OTUs responded to time lag. Two bacterioplankton communities from dissimilar ponds were inoculated into one of the natural media with a time lag of 0, 12, 36 or 60 h, giving advantage in time to the foreign community . All three time lags resulted in priority effects, as the final community composition of these treatments differed significantly from that of the treatment with no time lag, but compositional shifts were not linear to inoculation time lag. The responses of individual OTUs to time lag were highly diverse and not predictable based on their immigration history or relative abundance in the inocula or control. The observed impact and complexity of priority effects in multispecies systems emphasize the importance of this process in structuring both natural and industrial bacterial communities.     

Control of bloom in a tropical lake: grazing efficiency of some herbivorous fishes

Using a limnocorral study, Microcystis grazing by silver carp, bighead and tilapia was examined in an eutrophic shallow lake permanently infested with a Microcystis bloom. Twelve fingerlings of each fish species were introduced into the limnocorrals placed in the lake, in triplicate, and their grazing effects were examined over a period until the limnocorral bloom declined substantially and remained unchanged. Introduction of fish, regardless of species, into the limnocorrals resulted in sharp reductions (60–93%) of the initial Microcystis population on day 3 or 7. Maximum gross and net clearance of Microcystis were by silver carp followed by bighead and tilapia. Short-term high retention of Microcystis by tilapia indicated that the clearance effect did not last long due to a high rate of defaecation of undigested Microcystis . Nearly 6–180% nutrient enrichment of the limnocorrals was attributable to the defaecation of test fishes suggesting their ichthyoeutrophication potentials, which were in the following order: tilapia>bighead>silver carp. It is concluded that silver carp is most suitable for clearing Microcystis in the lake because of its minimal ichthyoeutrophication effect.     

铜绿微囊藻对紫外辐射的生理代谢响应

采用紫外(UV)滤膜过滤日光UV以及紫外灯添加UV的方法,研究了UV辐射对铜绿微囊藻Microcystis aeruginosa单细胞藻株PCC7806和群体藻XW01生长及生理代谢的影响。结果显示,在室内条件下低剂量UV辐射可促进群体微囊藻XW01生长;室外条件下与滤除了UV的光照相比,含有UV的完全日光更有利于微囊藻生长;而相同的UV辐射强度均导致单细胞株死亡,群体株显示了较强的UV抗性;日光中的UV可促进XW01合成抗氧化相关的超氧化物歧化酶(SOD)和过氧化氢酶(CAT)、促进胞外多糖的产生并形成较大的群体、促进UV屏障物质类菌孢素氨基酸(MAAs)和伪枝藻素(Scy)积累。这些生理代谢的改变,消除了阳光辐射中UV对微囊藻的伤害。研究的结果提示,自然条件下阳光中的UV有助于群体微囊藻生长。     

Pseudomonas fluorescens enhances resistance and natural enemy population in rice plants against leaffolder pest

Plant growth promoting fluorescent pseudomonad strains Pf1, TDK1 and PY15 were evaluated for their efficacy against leaffolder ( Cnaphalocrocis medinalis ) pest in rice plants under field conditions individually and in combinations. Application of mixtures of Pseudomonas fluorescens strains Pf1, TDK1 and PY15 significantly reduced the leaffolder damage in rice plants compared with untreated control. Interestingly, natural enemy population in plots treated with P. fluorescens was greater than the chemical and untreated controls. Further, support for these results was gathered by assaying activities of polyphenol oxidase (PPO) and lipoxygenase (LOX) under glasshouse conditions. The results showed the higher activity of PPO and LOX in plants treated with P. fluorescens mixtures (Pf1 + TDK1 + PY15) than the plants treated with individual strains, chemical and untreated controls. Further, fluorescent pseudomonad mixtures increased the rice yield compared with individual strain and non-bacterized treatments. The present study reveals that in addition to plant growth promotion, plant growth-promoting rhizobacterial (PGPR) strains-mediated induction of PPO and LOX in rice plants could be involved in enhanced natural enemy populations and resistance mechanisms against leaffolder attack.