Feeding behavior of Brevicoryne brassicae in resistant and susceptible collard greens genotypes: interactions among morphological and chemical factors

The cabbage aphid, Brevicoryne brassicae (L.) (Hemiptera: Aphididae), is distributed throughout the tropical and subtropical areas of the world. The main crops attacked by B. brassicae are cabbage, collard greens, broccoli, Brussels sprouts, and cauliflower. To survive the attack of pest insects, plants have evolved various resistance mechanisms that may affect pest feeding behavior. The use of electronic monitoring through EPG (electrical penetration graph) can help characterize and distinguish the resistance mechanisms involved. This study evaluated the feeding behavior of B. brassicae in eight genotypes of collard greens, Brassica oleraceae L. var. acephala (Brassicaceae), exhibiting antixenosis and/or antibiosis resistance to this insect. Possible correlations were established between the glucosinolate levels, the hardness, and the epicuticular wax on the leaves vs. aphid feeding behavior. On the genotypes 22V, 5E, and 27VA, for which many ‘potential drop’ waves were performed, aphid development was slower, indicating antixenosis as resistance type. Aphids on the genotypes 22V and 24X required more time until accessing the phloem, also suggesting antixenosis as resistance category. Genotypes 22V and PE had hard leaves, which also points at antixenosis. Genotypes 20T and HS had higher total wax and wax mg⁻¹. Feeding parameters on ARI and 24X were similar to those observed on HS; antibiosis is likely to be the predominant resistance category of this germplasm. Because HS was considered as a susceptible standard genotype in this study, a higher gluconapin amount indicates that this compound does not influence cabbage aphid feeding behavior. The present study confirms that analysis of the physical and chemical aspects of collard greens genotypes by the EPG technique can provide a useful approach for the study of plant resistance to cabbage aphids.     

An investigation of the resistance to cabbage aphid in brassica species

Fifteen Brassica species and subspecies, as well as accessions of Arabidopsis thaliana and Eruca sativa, were evaluated for their resistance to the cabbage aphid, Brevicoryne brassicae, in the field and laboratory at Horticulture Research International (HRI) Wellesbourne in 1992. In the laboratory, aphid performance was measured in terms of pre-reproductive period, reproductive period, population increase and insect survival. Using these parameters it was possible to identify brassicas possessing antibiosis resistance. In the field B. brassicae attack was assessed in terms of numbers of insects and it was possible to determine aphid preferences for alighting on different species. High levels of antixenosis and antibiosis resistance were discovered in accessions of Brassica fruticulosa and B. spinescens, in a Brassica juncea breeding line and in Eruca sativa. Partial resistance was found in several other Brassica species. The potential of these various sources as donors of resistance genes to be bred into cultivated brassicas is discussed.     

The influence of cultivar and cultivar-aphid odours on the olfactory response of the parasitoid Aphidius colemani

The olfactory response of the parasitoid Aphidius colemani (Viereck) (Hymenoptera: Braconidae) to odours in a tritrophic system involving three cultivars of common cabbage, Brassica oleracea var capitata, characterized by different levels of susceptibility to Myzus persicae (Sulzer) (Hemiptera: Aphididae) was studied in a four‐way olfactometer. Odours influenced A. colemani response in the olfactometer to varying degrees. The magnitude of parasitoid response to odours of uninfested cabbage depended on cultivar, with Derby Day [green‐leaved, susceptible to M. persicae and the crucifer specialist, Brevicoryne brassicae (Linnaeus) (Hemiptera: Aphididae)] and Minicole (green‐leaved, partially resistant with known antibiosis factors for B. brassicae) preferred over Ruby Ball (red‐leaved with antixenosis factors for M. persicae and B. brassicae). The odour of the cabbage cultivar on which the parasitoid had been reared was preferred over the other cultivars. However, when provided with a choice between odours of infested plants, parasitoids did not show a significant preference for the cultivar on which they were reared. Results from the study show that parasitioids differentiated between odour of the three cultivars in dependence of their rearing history when the plant is uninfested.     

Field and glasshouse studies on components of resistance to root fly attack in swedes

Field trials in two years indicated that egg-laying antixenosis was the major component of resistance in swede cvs Angus and Melfort against root flies (Delia floralis and D. radicum). Between four and eight times as many D. floralis eggs were laid on the susceptible cv. Doon Major as on the resistant cv. Angus. The degree of antixenosis effective against D. radicum was more variable in 1985, due to low numbers of adult flies, but in 1986, when adult fly numbers were higher, D. radicum laid 34 times as many eggs on susceptible cv. Sator Øtofte as on Angus. Root antibiosis against larval feeding is a second but less important component of resistance against D. floralis. In egg inoculation tests, 1.3-2.0 times more D. floralis pupae developed on Doon Major than on Angus or Melfort. Larval feeding on resistant cultivars was restricted to surface root tissue only, compared with deeper tunnelling on susceptible cultivars. Tolerance to root damage may also occur under certain conditions. Although increased dry matter content was significantly correlated with root antibiosis, its effect on larval feeding was apparently not due to increased tissue hardness. It seems likely that chemical differences in the outer root tissues influence larval feeding and development.     

Effect of cultivars with varying levels of resistance to aphids on development time,sex ratio,size and longevity of the parasitoid <Emphasis Type="Italic">Aphidius colemani</Emphasis>

The influence of cultivars of common cabbage, Brassica oleracea var. capitata with varying levels of resistance to Brevicoryne brassicae (L.) and Myzus persicae (Sulzer) on key biological characteristics of Aphidius colemani (Viereck) was investigated under laboratory conditions. The total development time for female parasitoids reared on M. persicae did not differ significantly between Minicole (green-leaved, partially resistant with antibiosis factors for B. brassicae) and Derby Day (green-leaved, susceptible to both aphid species); but development was significantly faster (ca 10%) on Ruby Ball (red-leaved, partially resistant with antixenosis factors for B. brassicae). Total development time for females reared on B. brassicae was slightly shorter on Ruby Ball than on Minicole. Males reared on M. persicae developed into adults significantly faster (ca 10%) on Ruby Ball than on Minicole. However, when B. brassicae was the host, no significant variations in development time were observed. Sex ratios, size and longevity of both male and female parasitoids on either host were not significantly influenced by cultivar. The results are discussed in relation to the compatible utilisation of host-plant resistance and biological control in the integrated management of aphids.     

Assessing Common Bean Cultivars for Resistance to the Soybean Looper <Emphasis Type="Italic">Chrysodeixis includens</Emphasis> (Lepidoptera: Noctuidae)

The soybean looper Chrysodeixis includens (Walker) (Lepidoptera: Noctuidae) is known as an important pest of leguminous plants worldwide. In Brazil, this pest species is gaining importance to producers of the common bean Phaseolus vulgaris L. (Fabaceae) because it limits field production of the crop. Chemical control is still the primary method of insect control. However, due to the possible harmful effects of pesticides to humans and the environment, alternative and less aggressive practices are being investigated. For this reason, the use of resistant plant genotypes represents a valuable tool in insect control. This study evaluated the biological aspects of larvae of C. includens confined to 14 bean genotypes under laboratory conditions (26 ± 2°C; 65 ± 10% RH; photoperiod of 14 h L:10 h D). The duration of the instars, total duration of the larval phase, consumption while in the larval phase, weight of the fifth instar larvae, larval viability, duration of the pre-pupal and pupal phases, pupal weight, pupal viability, pupal deformity, caterpillar-to-adult cycle, duration of the pre-oviposition and oviposition periods, and total number of viable eggs per female were evaluated. The genotypes “IAC Boreal,” “IAC Harmonia,” and “IAC Formoso” expressed antibiosis, prolonging the caterpillar-to-adult cycle and reducing the larval viability; however, each of these genotypes also experienced high leaf consumption. “IAC Jabola” expressed moderate levels of antibiosis and/or antixenosis (feeding), while the genotype “BRS Horizonte” expressed antixenosis (feeding). The data obtained with IAC Boreal, IAC Harmonia, IAC Formoso, IAC Jabola, and BRS Horizonte are promising and may help with the improvement of programs aimed at managing C. includens damage to this leguminous agricultural crop.     

Incorporation of Novel <Emphasis Type="Italic">Ceutorhynchus obstrictus</Emphasis>-resistant Canola Genotypes Into Mixed Cropping Strategies: Evidence for Associational Resistance

Canola genotypes resistant to the cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham) (Coleoptera: Curculionidae), have recently been developed through introgression of Sinapis alba L. to Brassica napus L. Several lines express antixenosis and antibiosis resistance and have been shown to be less attractive to weevils in visual and olfactory behavioral bioassays. This paper details a small-plot study that assessed the effects on distribution dynamics of weevil adults and larvae of interspersing susceptible among resistant genotypes relative to monocultures over two growing seasons. Results indicate that mixes reduced weevil numbers and oviposition in pods of susceptible genotypes. These results are consistent with associational resistance.     

Good mothers,bad mothers,and the nature of resistance to herbivory in <Emphasis Type="Italic">Solidago altissima</Emphasis>

Evidence of poor correspondence between an insect herbivore’s oviposition preferences and the performance of its offspring has generally been attributed either to maladaptive behavior of the insect mother or inadequate measurement by the researcher. In contrast, we hypothesize that many cases of “bad mothers” in herbivores may be a byproduct of the hierarchical way natural selection works on resistance in host plants. Epistatic selection on the components of resistance (i.e., antixenosis and antibiosis) may generate negative genetic correlations between the resistance components, which could counteract the efforts of herbivores to oviposit on the best hosts for the performance of their offspring. In common garden and greenhouse experiments, we measured aspects of antixenosis and antibiosis resistance in 26 genets of tall goldenrod, Solidago altissima, against two common herbivores: the gall-inducing fly Eurosta solidaginis and the spittlebug Philaenus spumarius. Goldenrod antixenosis and antibiosis were positively correlated against E. solidaginis and negatively correlated against P. spumarius. Analogously, population-wide preference–performance correlations were positive for the gall flies and negative for the spittlebugs. Several natural history differences between the two insects could make gall flies better mothers, including better synchrony of the phenologies of the flies and the host plant, the much narrower host range of the gall flies than the spittlebugs, and the more sedentary lifestyle of the gall fly larvae than the spittlebug nymphs. If these results are typical in nature, then negative genetic correlations in antixenosis and antibiosis in plants may often result in zero or negative population-wide correlations between preference and performance in herbivores, and thus may be an important reason why herbivorous insects often appear to be bad mothers.     

Reevaluating the conceptual framework for applied research on host‐plant resistance

Applied research on host‐plant resistance to arthropod pests has been guided over the past 60 years by a framework originally developed by Reginald Painter in his 1951 book, Insect Resistance in Crop Plants. Painter divided the “phenomena” of resistance into three “mechanisms,” nonpreference (later renamed antixenosis), antibiosis, and tolerance. The weaknesses of this framework are discussed. In particular, this trichotomous framework does not encompass all known mechanisms of resistance, and the antixenosis and antibiosis categories are ambiguous and inseparable in practice. These features have perhaps led to a simplistic approach to understanding arthropod resistance in crop plants. A dichotomous scheme is proposed as a replacement, with a major division between resistance (plant traits that limit injury to the plant) and tolerance (plant traits that reduce amount of yield loss per unit injury), and the resistance category subdivided into constitutive/inducible and direct/indirect subcategories. The most important benefits of adopting this dichotomous scheme are to more closely align the basic and applied literatures on plant resistance and to encourage a more mechanistic approach to studying plant resistance in crop plants. A more mechanistic approach will be needed to develop novel approaches for integrating plant resistance into pest management programs.     

Antibiosis resistance in soybean plant introductions to Dectes texanus (Coleoptera: Cerambycidae)

No soybean cultivars exhibit resistance to larval damage by the cerambycid, Dectes texanus LeConte, in the United States. Selected soybean varieties and plant introductions (PIs) in maturity groups VI to VIII from the U.S. Department of Agriculture National Soybean Germplasm Collection were evaluated for D. texanus resistance in a series of field and laboratory experiments from 2005 through 2008. In field cage experiments, the numbers of oviposition punctures (OPs) per plant were determined as indicators of oviposition antixenosis and the ratio of OPs per live D. texanus larvae (OP/Lv) served as an indicator of plant antibiosis to larvae. A D. texanus-susceptible variety treated with the systemic insecticide fipronil was used as a positive antibiosis control. Plant introduction PI165673 had the highest OP/Lv ratio, indicating that even if oviposition was successful, an antibiosis factor in PI165673 significantly reduced egg hatch and the resulting number of live D. texanus larvae. Factorial analyses indicated that maturity group is not a significant factor in the expression of resistance. Thus, PI165673 appears to be a potential source of resistance to D. texanus. In related field studies, the preferred D. texanus oviposition site was localized to leaf petioles in the upper four or five nodes of the plant canopy. Histomorphological analyses of petiole cross-sections of plant introductions PI171451, PI165676, and PI165673 indicated that leaf petiole morphology may be related to reduced D. texanus oviposition on petioles of PI171451 and PI165676, but that resistance in PI165673 is independent of petiole morphology.     

Protocol for assessing soybean antixenosis to Heliothis virescens

Larvae of Heliothis virescens (Fabricius) (Lepidoptera: Noctuidae) often infest soybean crops, Glycine max (L.) (Fabaceae), causing significant yield losses in important soybean-producing regions. The use of soybean varieties resistant to lepidopteran larvae is a major approach in soybean integrated pest management. However, standardization and optimization of bioassays that are used to screen genotypes for insect resistance are essential for high-throughput phenotyping. Methodologies for screening were assessed to determine the most effective method for discriminating levels of antixenosis to H. virescens in soybean plants. Feeding and oviposition preference assays were performed to determine optimal densities of larvae and adults, and optimal plant structures and growth stages for conducting assays. In addition, trichome densities, and fiber and lignin contents were quantified in plant structures of soybean cultivars differing in resistance. Resistance levels of cultivars were best differentiated using nine neonate larvae and two 6-day-old larvae, and by using young leaves of plants at the vegetative stage. This was likely due to the more pronounced differences in lignin and fiber contents in young leaves of vegetative-stage plants. Density of adult pairs, plant structure, and growth stage did not affect ability to distinguish differences in oviposition preference by H. virescens. Higher numbers of eggs were found on the leaves, which were the plant structures that exhibited the lowest trichome densities. The protocol developed in this work will benefit future evaluations of soybean genotypes for antixenosis against H. virescens.     

Mechanisms of resistance to Helicoverpa armigera and introgression of resistance genes into F1 hybrids in chickpea

The noctuid pod borer, Helicoverpa armigera is a major pest of chickpea, and host plant resistance is an important component for managing this pest. We evaluated a set of diverse chickpea genotypes with different levels of resistance to H. armigera, and their F₁ hybrids for oviposition non-preference, antibiosis, and tolerance components of resistance under uniform insect infestation under greenhouse/laboratory conditions. The genotypes ICC 12476, ICC 12477, ICC 12478, ICC 12479, and ICC 506EB were non-preferred for oviposition under no-choice, dual-choice, and multi-choice conditions, and also suffered lower leaf damage in no-choice tests as compared to the susceptible check, ICCC 37. Antibiosis expressed in terms of low larval weights was observed in insects reared on ICC 12476, ICC 12478, and ICC 506EB. Weight gain by the third-instars was also low on ICC 12476, ICC 12477, ICC 12478, ICC 12479, and ICC 506EB at the podding stage. Non-preference for oviposition and antibiosis (poor larval growth) were also expressed in hybrids based on ICC 12477, ICC 12476, ICC 12478, ICC 12479, and ICC 506EB as compared to the hybrids based on the susceptible check, ICCC 37, indicating that oviposition non-preference and antibiosis in the F₁ hybrids is influenced by the parent genotype. Loss in grain yield was lower in ICC 12477, ICC 12478, ICC 12479, and ICC 506EB compared to that on ICCC 37. The genotypes ICC 12477, ICC 12478, ICC 12479, and ICC 506EB showing antixenosis, antibiosis, and tolerance mechanism of resistance to H. armigera can be used for developing chickpea cultivars for resistance to this pest.     

Effects of different host plants and rearing atmosphere on life cycle of large white cabbage butterfly,Pieris brassicae (Linnaeus)

We studied the effect of different host plants and rearing atmosphere on life cycle of cabbage butterfly, Pieris brassicae. Insects were reared in the field (fluctuating weather) as well as in the laboratory (constant rearing conditions) on four host plants, viz. cabbage, cauliflower, knol-khol and broccoli. Significant differences were not found in the incubation and pupal period of butterfly reared on different host plants. However, larval period was found to be significantly lower on cabbage followed by knol-khol and highest on broccoli. Therefore, the developmental period (from eggs to adult) was reasonably lower on cabbage. Furthermore, inverse relationship was found in the body weight of butterfly and developmental period, where weight of full grown caterpillar and pupae was significantly higher on cauliflower and lowest on cabbage. Besides, significant differences were not found in the body weight of P. brassicae caterpillar reared under field and in the laboratory. Nevertheless, pupal weight of butterfly was significantly higher under field conditions than the laboratory conditions. Overall, the development of P. brassicae was much faster on cabbage than other hosts; but its body weight was considerably higher on cauliflower.     

Detection and analysis of QTLs for resistance to the brown planthopper, Nilaparvata lugens, in a doubled-haploid rice population

 We used a mapping population of 131 doubled-haploid lines, produced from a cross between an improved indica rice variety (IR64) and a traditional japonica variety (Azucena), to detect quantitative trait loci (QTLs) for resistance to the brown planthopper (BPH), Nilaparvata lugens. We evaluated the parents and mapping population with six tests that measure varying combinations of the three basic mechanisms of insect host plant resistance, i.e., antixenosis, antibiosis, and tolerance. To factor-out the effect of the major resistance gene Bph1 from IR64, the screening was done with two BPH populations from Luzon Island, The Philippines, that are almost completely adapted to this gene. A total of seven QTLs associated with resistance were identified, located on 6 of the 12 rice chromosomes. Individual QTLs accounted for between 5.1 and 16.6% of the phenotypic variance. Two QTLs were predominantly associated with a single resistance mechanism: one with antixenosis and one with tolerance. Most of the QTLs were derived from IR64, which has been shown to have a relatively durable level of moderate resistance under field conditions. The results of this study should be useful in transferring this resistance to additional rice varieties. Received: 10 May 1998 / Accepted: 4 June 1998     

Asymmetric plant-mediated cross-effects between a herbivorous insect and a phytopathogenic fungus

• 1 Cross‐effects between a herbivorous insect and a phytopathogenic fungus on their common host plant were examined. Specifically, we addressed the questions whether (i) infection of Chinese cabbage leaves by the fungus Alternaria brassicae affects the development and host selection behaviour of the leaf beetle Phaedon cochleariae and (ii) whether herbivory influences host suitability of Chinese cabbage for A. brassicae.
• 2 Feeding on fungus‐infected leaves prolonged larval development and reduced pupal weight of P. cochleariae. Adult beetles avoided feeding and egg deposition on fungus‐infected leaves. In contrast to these local effects, no systemic effect of phytopathogenic infection on the herbivore was detected.
• 3 Herbivory did not influence fungal growth neither locally nor systemically.
• 4 Thus, our results demonstrate an asymmetric relationship between herbivore and fungus. Whereas herbivory had no visible impact on fungal growth, fungal infection of the plant induced local resistance against P. cochleariae.

     

Ovipositional preferences and larval survival of annual bluegrass weevil,Listronotus maculicollis,on Poa annua and selected bentgrasses (Agrostis spp.)

The annual bluegrass weevil (ABW), Listronotus maculicollis Kirby (Coleoptera: Curculionidae), is a serious and expanding pest of short‐cut turfgrass on golf courses in eastern North America. Increasing problems with the development of insecticide resistance in this pest highlights the need for more sustainable management approaches. Plant resistance is one of the most promising alternative strategies. Bentgrasses are the dominant grass species on golf course fairways, tees, and putting greens in the areas affected by ABW. But Poa annua L. (Poaceae), a highly invasive weed, often constitutes a large percentage of turf stands in short‐mown golf courses and is thought to be particularly susceptible to ABW. We studied resistance to ABW in four cultivars of creeping bentgrass, Agrostis stolonifera L., and two cultivars each of colonial bentgrass, Agrostis capillaris L., and velvet bentgrass, Agrostis canina L. (Poaceae), in comparison with P. annua by addressing the three major components of resistance: antixenosis (adult ovipositional and feeding preferences), antibiosis (larval survival and growth), and grass tolerance (grass damage). Our findings suggest that antixenosis/non‐preference is at least partially involved in bentgrass resistance to ABW. Even though oviposition was observed in all tested grasses, females laid significantly fewer eggs in Agrostis spp. than in P. annua. Compared to P. annua, Agrostis spp. were also less suitable for larval development with lower numbers of ABW immatures recovered and larvae weighing less and being less advanced in development. Resistance levels to ABW larvae varied significantly among Agrostis spp. and cultivars. Agrostis canina was least preferred by females for oviposition and A. stolonifera was the least suitable for larval survival and development. Agrostis spp., especially A. stolonifera, were more tolerant to ABW feeding than P. annua. Our findings suggest that reduction in P. annua and replacement with Agrostis spp., especially A. stolonifera, wherever feasible should be integral to more sustainable approaches to ABW management.     

Comparative headspace analysis of cabbage plants damaged by two species ofPieris caterpillars: consequences for in-flight host location byCotesia parasitoids

Headspace composition, collected from intact cabbage plants and cabbage plants infested with eitherPieris brassicae L. orP. rapae L. (Lepidoptera: Pieridae) first instar larvae, was determined by GC-MS. Twenty-one volatiles were identified in the headspace of intact plants. Twenty-two volatiles were identified in the headspace of plants infested byP. brassicae larvae, 2 of which, Z-3-hexenyl butyrate and Z-3-hexenyl isovalerate, were not detected in the headspace of either intact orP. rapae damaged plants. In the headspace of the latter, 21 compounds were identified, all of which which were also produced by intact plants. No significant quantitative differences were found between headspace composition of the plants damaged by one or the other caterpillar species. Major differences between intact and caterpillar-damaged plants in contribution to the headspace profile were revealed for hexyl acetate, Z-3-hexenyl acetate, myrcene, sabinene and 1,8-cineole. The larval endoparasitoidCotesia glomerata L. was attracted by the volatiles emanating fromB. oleracea damaged byP. brassicae first instar larvae.C. rubecula L., a specialized larval endoparasitoids ofP. rapae, was attracted by the volatiles released from theB. oleracea-P. rapae plant-host complex. This shows that cabbage plants kept under the conditions of headspace collection produce attractive volatiles for both parasitoids.     

Common cabbage resistance mechanisms against the diamondback moth: still an open book?

Despite the extensive research on host plant resistance of Brassica oleracea L. var. capitata (common cabbage) against Plutella xylostella L. (diamondback moth), knowledge of the resistance mechanisms involved remains incomplete. While robust evidence exists for the physical (antixenosis) mechanisms believed to cause resistance in glossy-leaved breeding lines, the one mechanism proposed in the literature to date (polar leaf extracts causing antibiosis) at best only explains a part of the resistance observed in two partially-resistant normal-bloom breeding lines in the field and the evidence is suggestive rather than direct.
Evidence from other host plant resistance studies involving P. xylostella is discussed and the other possible mechanisms are considered with the aim of providing a context for further investigations on host plant resistance mechanisms against P. xylostella.     

A comparative study of the egg-laying behaviour and larval development of Pieris rapae L. and P. brassicae L. on the same host plants

Summary Pieris rapae and P. brassicae feed on the same host plants and have synchronized seasons. P. brassicae, whose larvae are twice the size of P. rapae, lays eggs in clusters of 40–100 eggs whereas P. rapae lays single eggs. In this paper we examine how egg clustering may be advantageous for P. brassicae. The larval development of each species was studied, and found not to differ significantly. P. brassicae larvae were observed to migrate from their host plant after defoliating it. A comparison of the efficiency of host plant utilization by the two pierid species was undertaken by measuring the effect of larval feeding on the growth of their host plants (kale and brussel sprouts). The results show that egg clustering is advantageous for larval fitness in terms of host resource exploitation, and we suggest that P. brassicae is adapted for ovipositing on clumped vegetation, while P. rapae is selected for exploiting isolated plants.