Plant Volatiles and Oviposition Behavior in the Selection of Barley Cultivars by Wheat Stem Sawfly (Hymenoptera: Cephidae).

Wheat stem sawfly, [Cephus cinctus (Hymenoptera: Cephidae)], females display complex behaviors for host selection and oviposition. Susceptible hollow stem wheat (Triticum aestivum L.) cultivars release a greater amount of attractive compound, (Z)-3-hexenyl acetate and receive a greater number of eggs compared to resistant solid stem wheat cultivars. However, barley (Hordeum vulgare L.) is becoming a more common host for C. cinctus in Montana. Therefore, how do host selection and oviposition behaviors on barley cultivars compare to what happens when encountering wheat cultivars? To answer this question, we carried out greenhouse experiments using two barley cultivars: 'Hockett' and 'Craft'. Between these cultivars at Zadoks stages 34 and 49, we compared host selection decisions using a Y-tube olfactometer, compared oviposition behaviors on stems, and counted the number of eggs inside individual stems. In Y-tube bioassays, we found a greater number of C. cinctus females were attracted to the airstream passing over 'Hockett' than 'Craft' barley cultivars. Although the frequencies of oviposition behaviors were similar between these cultivars, the number of eggs was greater in 'Hockett'. Volatile profiles indicated that the amount of linalool was greater in the airstream from 'Craft' than in 'Hockett' at Zadoks 34 while the amount of (Z)-3-hexenyl acetate was greater in airstream from 'Hockett' at both Zadoks 34 and 49. These results suggest that volatiles of barley plants influenced host selection behavior of ovipositing C. cinctus females, while other discriminating behaviors do not differ between cultivars.

Multiple decrement life tables of Cephus cinctus Norton (Hymenoptera: Cephidae) across a set of barley cultivars: The importance of plant defense versus cannibalism.

Accurately estimating cause-specific mortality for immature insect herbivores is usually difficult. The insects are exposed to abiotic and biotic mortality factors, causing cadavers to simply disappear before cause of mortality can be recorded. Also, insect herbivores are often highly mobile on hosts, making it difficult to follow patterns for individuals through time. In contrast, the wheat stem sawfly, Cephus cinctus Norton, spends its entire egg, larval, and pupal period inside a host stem. Therefore, with periodic sampling stage-specific causes of mortality can be ascertained. Consequently, we examined C. cinctus mortality in eight barley, Hordeum vulgare L., cultivars in two locations in Montana from 2016 to 2018 by collecting stem samples from stem elongation to crop maturity at weekly intervals, and collecting overwintered barley stubs the following spring and summer from the same plots. If larvae were present, we examined larval status-dead or alive-and categorized dead individuals into one of 5 mortality categories: plant defense, cannibalism, parasitism, pathogens, and unknown factors. We used multiple decrement life tables to estimate cause-specific mortality and irreplaceable mortality (the proportion of mortality from a given cause that cannot be replaced by other causes of mortality). Plant defense (antibiosis) caused 85.7 ± 3.6%, cannibalism (governed by antixenosis) caused 70.1 ± 7.6%, parasitism caused 13.8 ± 5.9%, unknown factors caused 38.5 ± 7.6%, and pathogens caused 14.7 ± 8.5% mortality in the presence of all causes of mortality. Similarly, irreplaceable mortality due to plant defense was 22.3 ± 6.4%, cannibalism was 29.1± 4.2%, unknown factors was 6.2 ± 1.8%, pathogens was 0.9 ± 0.5%, and parasitism was 1. 5 ± 0. 6%. Antibiosis traits primarily killed newly emerged larvae, while other traits supported more favorable oviposition decisions by females, increasing mortality by obligate cannibalism. Our results suggest that breeding barley for resistance to C. cinctus targeting both categories of traits (antibiosis and antixenosis) is a highly valuable tactic for management of this important pest.

Antixenosis, Antibiosis, and Potential Yield Compensatory Response in Barley Cultivars Exposed to Wheat Stem Sawfly (Hymenoptera: Cephidae) Under Field Conditions.

Wheat stem sawfly, Cephus cinctus Norton, is an economically serious pest of cereals grown in North America. Barley cultivars were previously planted as resistant crops in rotations to manage C. cinctus, but due to increasing levels of injury to this crop, this is no longer a valid management tactic in Montana. Therefore, we aimed to understand antixenosis (behavioral preference), antibiosis (mortality), and potential yield compensation (increased productivity in response to stem injuries) in barley exposed to C. cinctus. We examined these traits in eight barley cultivars. Antixenosis was assessed by counting number of eggs per stem and antibiosis was assessed by counting infested stems, dead larvae, and stems cut by mature larvae. Potential yield compensation was evaluated by comparing grain yield from three categories of stem infestation: 1) uninfested, 2) infested with dead larva, and 3) infested cut by mature larva at crop maturity. We found the greatest number of eggs per infested stem (1.80 ± 0.04), the highest proportion of infested stems (0.63 ± 0.01), and the highest proportion of cut stems (0.33 ± 0.01) in 'Hockett'. Seven out of eight cultivars had greater grain weight for infested stems than for uninfested stems. These cultivars may have compensatory responses to larval feeding injury. Overall, these barley cultivars contain varying levels of antixenosis, antibiosis, and differing levels of yield compensation. Our results provide foundational knowledge on barley traits that will provide a framework to further develop C. cinctus resistant or tolerant barley cultivars.

Effect of Precipitation and Temperature on Larval Survival of Cephus cinctus (Hymenoptera: Cephidae) in Barley Cultivars.

Host plant traits strongly affect survivorship of insect herbivores, and host suitability is especially important for the wheat stem sawfly, Cephus cinctus Norton, which spends its entire egg, larval, and pupal periods in a single stem. Measuring larval survival inside stems from egg hatch through diapause-mediated dormancy is a potential measure of population size for the next year but is also useful in assessing effects of growing season precipitation and temperature. Larval growth is synchronized with host plant growth, and the larva cannot switch hosts. Thus, incorporating plant physiological time, as growing degree days (GDD), may yield a better prediction of larval survival. Therefore, we assessed wheat stem sawfly survival from early larval growth to the beginning of autumnal diapause in barley cultivars selected from across feed, forage, and two- or six-row malt groups. Field experiments were conducted in Gallatin and Chouteau counties, Montana, in 2016 and 2017. We used Kaplan-Meier estimation to assess larval survival among cultivars. We found that the survival of pre-diapause larvae was greatest in 'Hockett' (36.5%) and lowest in 'Celebration' (15.4%). Precipitation and temperature during the growing season affected temporal patterns for larval survival across study sites. Adjusting survivorship curves using site-specific GDD accumulation allowed cultivar-specific survivorship to be estimated more precisely for each site, despite differing environmental influences. Our findings suggest that measuring wheat stem sawfly survival across barley cultivars and standardizing by site-specific GDDs may provide better recommendations on barley cultivars that impede wheat stem sawfly population growth and reduce economic losses.

Characterization of Resistance to Cephus cinctus (Hymenoptera: Cephidae) in Barley Germplasm.

Most barley cultivars have some degree of resistance to the wheat stem sawfly (WSS), Cephus cinctus Norton (Hymenoptera: Cephidae). Damage caused by WSS is currently observed in fields of barley grown in the Northern Great Plains, but the impact of WSS damage among cultivars due to genetic differences within the barley germplasm is not known. Specifically, little is known about the mechanisms underlying WSS resistance in barley. We characterized WSS resistance in a subset of the spring barley CAP (Coordinated Agricultural Project) germplasm panel containing 193 current and historically important breeding lines from six North American breeding programs. Panel lines were grown in WSS infested fields for two consecutive years. Lines were characterized for stem solidness, stem cutting, WSS infestation (antixenosis), larval mortality (antibiosis), and parasitism (indirect plant defense). Variation in resistance to WSS in barley was compared to observations made for solid-stemmed resistant and hollow-stemmed susceptible wheat lines. Results indicate that both antibiosis and antixenosis are involved in the resistance of barley to the WSS, but antibiosis seems to be more prevalent. Almost all of the barley lines had greater larval mortality than the hollow-stemmed wheat lines, and only a few barley lines had mortality as low as that observed in the solid-stemmed wheat line. Since barley lines lack solid stems, it is apparent that barley has a different form of antibiosis. Our results provide information for use of barley in rotation to control the WSS and may provide a basis for identification of new approaches for improving WSS resistance in wheat.

Herbivore-mediated effects of glucosinolates on different natural enemies of a specialist aphid.

The cabbage aphid Brevicoryne brassicae is a specialist herbivore that sequesters glucosinolates from its host plant as a defense against its predators. It is unknown to what extent parasitoids are affected by this sequestration. We investigated herbivore-mediated effects of glucosinolates on the parasitoid wasp Diaeretiella rapae and the predator Episyrphus balteatus. We reared B. brassicae on three ecotypes of Arabidopsis thaliana that differ in glucosinolate content and on one genetically transformed line with modified concentrations of aliphatic glucosinolates. We tested aphid performance and the performance and behavior of both natural enemies. We correlated this with phloem and aphid glucosinolate concentrations and emission of volatiles. Brevicoryne brassicae performance correlated positively with concentrations of both aliphatic and indole glucosinolates in the phloem. Aphids selectively sequestered glucosinolates. Glucosinolate concentration in B. brassicae correlated negatively with performance of the predator, but positively with performance of the parasitoid, possibly because the aphids with the highest glucosinolate concentrations had a higher body weight. Both natural enemies showed a positive performance-preference correlation. The predator preferred the ecotype with the lowest emission of volatile glucosinolate breakdown products in each test combination, whereas the parasitoid wasp preferred the A. thaliana ecotype with the highest emission of these volatiles. The study shows that there are differential herbivore-mediated effects of glucosinolates on a predator and a parasitoid of a specialist aphid that selectively sequesters glucosinolates from its host plant.