Anne-Marja Nerg

Emission of Plutella xylostella-Induced Compounds from Cabbages Grown at Elevated CO2 and Orientation Behavior of the Natural Enemies

Several plant species defend themselves indirectly from herbivores by producing herbivore-induced volatile compounds that attract the natural enemies of herbivores. Here we tested the effects of elevated atmospheric CO2 (720 μmol mol−1) concentration on this indirect defense, physiological properties, and constitutive and induced emissions of white cabbage (Brassica oleracea ssp. capitata, cvs Lennox and Rinda). We monitored the orientation behavior of the generalist predator Podisus maculiventris (Heteroptera: Pentatomidae) and the specialist parasitoid Cotesia plutellae (Hymenoptera: Braconidae) to plants damaged by Plutella xylostella (Lepidoptera: Plutellidae) in the Y-tube olfactometer. Elevated CO2 levels did not affect stomatal densities but reduced specific leaf area and increased leaf thickness in cv Lennox. In addition to enhanced constitutive monoterpene emission, P. xylostella-damaged cabbages emitted homoterpene (E)-4,8-dimethyl-1,3,7-nonatriene, sesquiterpene (E,E)-α-farnesene, and (Z)-3-hexenyl acetate. Growth at elevated CO2 had no significant effect on the emissions expressed per leaf area, while minor reduction in the emission of homoterpene (E)-4,8-dimethyl-1,3,7-nonatriene and (E,E)-α-farnesene was observed at elevated CO2 in one of two experiments. The generalist predator P. maculiventris discriminated only between the odors of intact and P. xylostella-damaged cv Rinda plants grown at ambient CO2 concentration, preferring the odor of the damaged plants. The specialist parasitoid C. plutellae preferred the odor of damaged plants of both cultivars grown at ambient CO2 but did not detect damaged cv Lennox plants grown at elevated CO2. The results suggest that elevated atmospheric CO2 concentration could weaken the plant response induced by insect herbivore feeding and thereby lead to a disturbance of signaling to the third trophic level.

Application of methyl jasmonate reduces growth but increases chemical defence and resistance against Hylobius abietis in Scots pine seedlings

Scots pine (Pinus sylvestris L., Pinaceae) produces a terpenoid resin which consists of monoterpenes and resin acids that offer protection against herbivores and pathogen attacks. Methyl jasmonate (MJ) is a potential plant elicitor which induces a wide range of chemical and anatomical defence reactions in conifers and might be used to increase resistance against biotic damage. Different amounts of MJ (control, 10 mm, and 100 mm) were applied to Scots pine to examine the vigour, physiology, herbivory performance, and induction of secondary compound production in needles, bark, and xylem of 2‐year‐old Scots pine seedlings. Growth decreased significantly in both MJ treated plants, and photosynthesis decreased in the 100 mm MJ treated plants, when compared to 10 mm MJ or control plants. The large pine weevil (Hylobius abietis L.) (Coleoptera: Curculionidae) gnawed a significantly smaller area of stem bark in the 100 mm treated plants than in the control or 10 mm treated plants. The 100 mm MJ treatment increased the resin acid concentration in the needles and xylem but not in the bark. Furthermore, both MJ treatments increased the number of resin ducts in newly developing xylem. The changes in plant growth and chemical parameters after the MJ treatments indicate shifts in carbon allocation, but MJ also affects plant physiology and xylem development. Terpenoid resin production was tissue‐specific, but generally increased after MJ treatments, which means that this compound may offer potential protection of conifers against herbivores.

Plant volatile organic compounds (VOCs) in ozone (O3) polluted atmospheres: the ecological effects.

Tropospheric ozone (O3) is an important secondary air pollutant formed as a result of photochemical reactions between primary pollutants, such as nitrogen oxides (NOx), and volatile organic compounds (VOCs). O3 concentrations in the lower atmosphere (troposphere) are predicted to continue increasing as a result of anthropogenic activity, which will impact strongly on wild and cultivated plants. O3 affects photosynthesis and induces the development of visible foliar injuries, which are the result of genetically controlled programmed cell death. It also activates many plant defense responses, including the emission of phytogenic VOCs. Plant emitted VOCs play a role in many eco-physiological functions. Besides protecting the plant from abiotic stresses (high temperatures and oxidative stress) and biotic stressors (competing plants, micro- and macroorganisms), they drive multitrophic interactions between plants, herbivores and their natural enemies e.g., predators and parasitoids as well as interactions between plants (plant-to-plant communication). In addition, VOCs have an important role in atmospheric chemistry. They are O3 precursors, but at the same time are readily oxidized by O3, thus resulting in a series of new compounds that include secondary organic aerosols (SOAs). Here, we review the effects of O3 on plants and their VOC emissions. We also review the state of current knowledge on the effects of ozone on ecological interactions based on VOC signaling, and propose further research directions.

Effects of elevated carbon dioxide and ozone on volatile terpenoid emissions and multitrophic communication of transgenic insecticidal oilseed rape (Brassica napus)

Does transgenically incorporated insect resistance affect constitutive and herbivore-inducible terpenoid emissions and multitrophic communication under elevated atmospheric CO(2) or ozone (O(3))? This study aimed to clarify the possible interactions between allocation to direct defences (Bacillus thuringiensis (Bt) toxin production) and that to endogenous indirect defences under future climatic conditions. Terpenoid emissions were measured from vegetative-stage non-Bt and Bt Brassica napus grown in growth chambers under control or doubled CO(2), and control (filtered air) or 100 ppb O(3). The olfactometric orientation of Cotesia vestalis, an endoparasitoid of the herbivorous diamondback moth (Plutella xylostella), was assessed under the corresponding CO(2) and O(3) concentrations. The response of terpenoid emission to CO(2) or O(3) elevations was equivalent for Bt and non-Bt plants, but lower target herbivory reduced herbivore-inducible emissions from Bt plants. Elevated CO(2) increased emissions of most terpenoids, whereas O(3) reduced total terpenoid emissions. Cotesia vestalis orientated to host-damaged plants independent of plant type or CO(2) concentration. Under elevated O(3), host-damaged non-Bt plants attracted 75% of the parasitoids, but only 36.8% of parasitoids orientated to host-damaged Bt plants. Elevated O(3) has the potential to perturb specialized food-web communication in Bt crops.

Emissions of volatile organic compounds and leaf structural characteristics of European aspen (Populus tremula) grown under elevated ozone and temperature

Northern forest trees are challenged to adapt to changing climate, including global warming and increasing tropospheric ozone (O(3)) concentrations. Both elevated O(3) and temperature can cause significant changes in volatile organic compound (VOC) emissions as well as in leaf anatomy that can be related to adaptation or increased stress tolerance, or are signs of damage. Impacts of moderately elevated O(3) (1.3x ambient) and temperature (ambient + 1 degrees C), alone and in combination, on VOC emissions and leaf structure of two genotypes (2.2 and 5.2) of European aspen (Populus tremula L.) were studied in an open-field experiment in summer 2007. The impact of O(3) on measured variables was minor, but elevated temperature significantly increased emissions of total monoterpenes and green leaf volatiles. Genotypic differences in the responses to warming treatment were also observed. alpha-Pinene emission, which has been suggested to protect plants from elevated temperature, increased from genotype 5.2 only. Isoprene emission from genotype 2.2 decreased, whereas genotype 5.2 was able to retain high isoprene emission level also under elevated temperature. Elevated temperature also caused formation of thinner leaves, which was related to thinning of epidermis, palisade and spongy layers as well as reduced area of palisade cells. We consider aspen genotype 5.2 to have better potential for adaptation to increasing temperature because of thicker photosynthetic active palisade layer and higher isoprene and alpha-pinene emission levels compared to genotype 2.2. Our results show that even a moderate elevation in temperature is efficient enough to cause notable changes in VOC emissions and leaf structure of these aspen genotypes, possibly indicating the effort of the saplings to adapt to changing climate.

Potential for the Use of Exogenous Chemical Elicitors in Disease and Insect Pest Management of Conifer Seedling Production

Elicitors are compounds, which activate chemical defences in plants. Various biosynthetic pathways are acti- vated in treated plants depending on the compound used. The most intensively studied elicitor for manipulating defence pathways in plants is methyl jasmonate, which modifies e.g. the production of terpenoids, the main constituents of conifer oleoresin. Other commonly tested chemical elicitors are salicylic acid, methyl salicylate and benzothiadiazole, which af- fect production of phenolic compounds in plants. Both jasmonate-based and salicylate-based elicitors have been shown to have suppressive effects on fungal diseases and insect pests of plants. So far, knowledge regarding the efficiency of elici- tor treatments for enhancing pest and fungal disease resistance of conifer seedlings is very limited. We review current knowledge of the effect of these elicitor compounds on pest and disease resistance in plants, and we analyze the potential pros and cons of using elicitors for future pest management strategies in forest nurseries.

Constitutive and herbivore-inducible glucosinolate concentrations in oilseed rape (Brassica napus) leaves are not affected by Bt Cry1Ac insertion but change under elevated atmospheric CO2 and O3

Glucosinolates are plant secondary compounds involved in direct chemical defence by cruciferous plants against herbivores. The glucosinolate profile can be affected by abiotic and biotic environmental stimuli. We studied changes in glucosinolate patterns in leaves of non-transgenic oilseed rape (Brassica napus ssp. oleifera) under elevated atmospheric CO2 or ozone (O3) concentrations and compared them with those from transgenic for herbivore-resistance (Bacillus thuringiensis Cry1Ac endotoxin), to assess herbivory dynamics. Both elevated CO2 and O3 levels decreased indolic glucosinolate concentrations in transgenic and non-transgenic lines, whereas O3 specifically increased the concentration of an aromatic glucosinolate, 2-phenylethylglucosinolate. The herbivore-inducible indolic glucosinolate response was reduced in elevated O3 whereas elevated CO2 altered the induction dynamics of indolic and aliphatic glucosinolates. Herbivore-resistant Bt plants experienced minimal leaf damage after target herbivore Plutella xylostella feeding, but exhibited comparatively similar increase in glucosinolate concentrations after herbivory as non-transgenic plants, indicating that the endogenous glucosinolate defence was not severely compromised by transgenic modifications. The observed differences in constitutive and inducible glucosinolate concentrations of oilseed rape under elevated atmospheric CO2 and O3 might have implications for plant–herbivore interactions in Brassica crop-ecosystems in future climate scenarios.

Long-term effects of exogenous methyl jasmonate application on Scots pine (Pinus sylvestris) needle chemical defence and diprionid sawfly performance

Scots pine [Pinus sylvestris L. (Pinaceae)] trees with four different seed origins were exposed to exogenous applications of the elicitor, methyl jasmonate (MeJA), for three consecutive years. We studied the effects of MeJA on needle chemistry (including monoterpenes, sesquiterpenes, and tricyclic resin acids), plant growth, and the performance of two diprionid sawflies, the European pine sawfly (Neodiprion sertifer Geoffr.) and the common pine sawfly (Diprion pini L.) (both Hymenoptera: Diprionidae). In general, foliar MeJA application affected the whole range of needle secondary chemistry with significantly higher concentrations of two monoterpenes, β‐pinene and limonene, in particular. Furthermore, for some seed origins the growth rates of N. sertifer and D. pini larvae were lower on needles of MeJA‐treated plants with either high total terpene or high resin acid concentrations. However, inconsistencies in diprionid sawfly performance within each studied Scots pine origin suggest genetic variance in needle secondary chemistry. The differences between selected seed origins and notably variable responses to MeJA application imply that adaptation of the seed to new conditions may have had an impact on secondary chemistry and, thus, on insect performance. Finally, our results suggest that modification of Scots pine defence by a low‐concentration exogenous elicitor affects the production of terpenoids in the newly growing needles, leading to poorer pine sawfly performance in origins with high terpenoid content, while not harming the growth of Scots pine trees.

Host location behavior of Cotesia plutellae Kurdjumov (Hymenoptera: Braconidae) in ambient and moderately elevated ozone in field conditions

In field O(3)-enrichment experiments increased herbivore densities have been reported, which could be due to negatively affected host location behavior of natural enemies. We addressed the impact of doubling background O(3) on the host location of the parasitoid Cotesia plutellae by conducting 24-h trials in an open-air O(3)-fumigation system during two consecutive years. Two circles (radii 1.40 and 4.00 m) of Plutella xylostella-infested potted cabbage plants were placed in the O(3) and ambient plots. Female wasps were released into each plot from the center, and observed 5 times over a 24-h period to assess their host location capability. Thereafter, plants were kept in laboratory conditions until larvae pupation to determine parasitism rates. No significant differences were detected between ambient and O(3)-enriched environments either in the number of wasps found in the field, or in the percentages of parasitized larvae. This suggests that moderately elevated O(3) will not affect the behavior of this parasitoid.

Significance of Wood Terpenoids in the Resistance of Scots Pine Provenances Against the Old House Borer, Hylotrupes bajulus, and Brown-Rot Fungus, Coniophora puteana

We tested how terpenoid (i.e., monoterpenes and resin acids) composition and concentration in wood affects resistance against wood-borers and decaying fungi. Scots pine (Pinus sylvestris) wood from nine provenances having variable terpenoid profiles was studied against the old house borer, Hylotrupes bajulus, and the decay fungus, Coniophora puteana. Provenances represented a 1200-km N–S transect from Estonia to northern Finland, but they were all cultivated for 7 years in the same nursery field, in central Finland. Mean relative growth rate (MRGR) of small H. bajulus larvae positively correlated with the total monoterpene concentration of wood, and feeding was associated with high proportion of levopimaric+palustric acid in wood. Provenance did not affect the MRGR of small or big larvae, but big larvae consumed more wood and produced more frass on the northern Ylitornio trees than on the southern Rakvere and Ruokolahti trees. Low β-pinene and total monoterpene concentration and low β: α-pinene ratio in wood were all associated with a high number of eggs. The most northern Muonio provenance was the most favored as an oviposition site, differing significantly from Saaremaa, Tenhola, and Suomussalmi. Wood from Saaremaa, Tenhola, Ruokolahti, and Suomussalmi provenance was most resistant against decay fungus, differing significantly from that of Kinnula provenance. However, decay resistance was not clearly associated with the concentrations of wood terpenoids. These results suggest that monoterpene composition of wood affects resistance against wood-boring Cerambycid beetles, but resistance against wood-decaying fungi is not as clearly associated with wood terpenoids.