Resistance to attack by Brevicoryne brassicae among plants of Brussels sprouts

Because they remained almost uncolonized by the cabbage aphid (Brevicoryne brassicae (L.)) throughout the growing season, plants of Brussels sprouts were singled out in each of 4 years, from plots heavily infested with the aphid, as possibly being resistant to attack. Clones of these plants were established from cuttings and tested in a controlled environment by inoculation with B. brassicae and later, in the field, by natural infestation. The tests confirmed that some of the plants were resistant to the aphid, and the most resistant of those from the first year of the work proved at least as resistant as any subsequently found. The resistance was expressed as antibiosis, but in the field host non-preference was also shown by incoming winged aphids. The possibility that biotypes of B. brassicae might exist, to which the resistant sprout clones were not necessarily resistant, was investigated using B. brassicae collected from sprouts from each of several areas in England. Eight sprout clones, seven of which were known to be resistant, and the other susceptible, to B. brassicae from Wellesbourne were tested with these other B. brassicae. The results showed that biotypes of the aphid, with differing abilities to colonize respective sprout clones, existed in each area, and of the seven sprout clones resistant to the Wellesbourne aphid, only one appeared never to be fully susceptible to one or more of the other biotypes of B. brassicae.     

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.     

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.     

Glasshouse evaluation of some systemic fungicides for control of clubroot of brassicae

Of nine systemic fungicides screened as soil mixes against clubroot, only the precursors of methyl benzimidazol-2-ylcarbamate (MBC) or ethyl benzimidazol-2-ylcarbamate (EBC) gave promising results. Benomyl was the most effective, usually giving control at 250 mg/kg dry soil. Most fungicides were less effective against an isolate of Plasmodiophora brassicae from Brussels sprouts than against one from rape. Disease control was slightly better on cabbage than on a highly susceptible rape variety.     

Status and Perspectives of Clubroot Resistance Breeding in Crucifer Crops

Clubroot disease is a major threat to crops belonging to the Brassicaceae. It is controlled most effectively by the use of resistant cultivars. Plasmodiophora brassicae, the causal agent, shows a wide variation for pathogenicity, which can be displayed by using differential host sets. Except for Brassica juncea and B. carinata, resistant accessions can be found in all major crops. Most resistance sources are race-specific, despite some race-independent resistant accessions which can be found in B. oleracea. European field isolates from P. brassicae display great variation and show a tendency to overcome different resistance sources from either B. rapa or B. oleracea. At present, resistance genes from stubble turnips (B. rapa) are most effective and most widely used in resistance breeding of different Brassica crops. Resistance to P. brassicae from turnips was introduced into Chinese cabbage, oilseed rape, and B. oleracea. Although most turnips carry more than one resistance gene, the resistant cultivars from other crops received primarily a single, dominant resistance gene having a race-specific effect. Populations of P. brassicae that are compatible against most of the used resistance sources have been present in certain European areas for many decades. Such pathogen populations appeared in Japanese Chinese cabbage crops only a few years after the introduction of resistant cultivars. As the spread of virulent P. brassicae pathotypes seems to be slow, resistant cultivars are still a very effective method of control in many cropping areas. Mapping studies have revealed the presence of several clubroot-resistance genes in the Brassica A and C genomes; most of these genes are showing race specificity. Only in B. oleracea was one broad-spectrum locus detected. Two loci from the A genome confer resistance to more than one pathotype, but not to all isolates. Progress made in the determination of resistance loci should be used to formulate and introduce an improved differential set. Future efforts for breeding P. brassicae resistance will focus on durability by broadening the genetic basis of clubroot resistance by using either natural variation or transgenic strategies.     

Canola quality affects second (<Emphasis Type="Italic">Brevicoryne brassicae</Emphasis>) and third (<Emphasis Type="Italic">Diaeretiella rapae</Emphasis>) trophic levels

Biological control agents can be used as a complementary control measure that can be combined with resistant host plants to control pests. In this study, the effects of different canola cultivars (Karaj-1, Karaj-2, Karaj-3, Licord, Okapi, Opera, RGS₀₀₃, Sarigol, Talaye and Zarfam) on the performance and life table parameters of the cabbage aphid, Brevicoryne brassicae, and its parasitoid, Diaeretiella rapae, were determined under laboratory conditions. Total fecundity of the cabbage aphid differed with cultivar, with the highest value (59.41 nymphs per female) of this parameter observed on Opera and the lowest (1.67) observed on RGS₀₀₃. The highest and lowest intrinsic rates of increase (r) of the cabbage aphid were observed on Opera (0.331 day^?1) and RGS₀₀₃ (? 0.242 day^?1) cultivars, respectively, suggesting these to be the most susceptible and most resistant cultivars to this pest. However, because the aphid did not settle and feed well on RGS₀₀₃, it was not possible to determine demographic parameters for its parasitoid. Consequently, the Okapi cultivar, which was the most resistant cultivar to the cabbage aphid after RGS₀₀₃, was used in this study to assess the parasitoid wasp. The parasitoid’s intrinsic rate of increase (r) varied from 0.426 day^?1 on the susceptible cultivar (Opera) to 0.341 day^?1 on the resistant canola cultivar Okapi. Aphid performance decreased 93% on the resistant canola cultivar, while parasitoid performance decreased only 20% on the resistant cultivar compared to more susceptible cultivar.     

The roles of ascospores and conidia of Pyrenopeziza brassicae in light leaf spot epidemics on winter oilseed rape (Brassica napus) in the UK

Ascospores of Pyrenopeziza brassicae were produced in apothecia (cup‐shaped ascomata) on oilseed rape debris. The conidia, which were morphologically identical to the ascospores, were produced in acervular conidiomata was greater than for lesions caused by ascospores. In June 2000, on the ground under a crop with light on the surface of living oilseed rape tissues. Ascospores were more infective than conidia on oilseed rape leaves. The proportion of lesions caused by conidia located on leaf veins leaf spot, numbers of petioles with apothecia decreased with increasing distance into the crop from the edge of pathways. Air‐borne ascospores of P. brassicae were first collected above debris of oilseed rape affected with light leaf spot on 5 October 1998 and 18 September 1999,12 or 23 days, respectively, after the debris had been exposed outdoors. P. brassicae conidia were first observed on leaves of winter oilseed rape on 6 January 1999 and 15 February 2000, respectively, after plots had been inoculated with debris in November 1998 and October 1999. In 1991/92, numbers of ascospores above a naturally infected crop were small from January to April and increased in June and July. P. brassicae conidia were first observed in February and the percentage plants with leaves, stems or pods with light leaf spot increased greatly in May and June. In 1992/93, in a crop inoculated with debris, numbers of airborne ascospores were small from October to January and increased from April to June. P. brassicae conidia were first observed on leaves in late November and light leaf spot was seen on stems and pods in March and June 1993, respectively.     

Survival of Alternaria brassicae and Alternaria brassicicola on crop debris of oilseed rape and cabbage

Alternaria brassicae and A. brassicicola lesions present on infected leaves of oilseed rape and cabbage placed outdoors on soil produced viable spores for as long as leaf tissues remained intact. For oilseed rape this was up to 8 wk and for cabbage up to 12 wk. On leaves exposed in November and January spore concentrations decreased with time but on leaves exposed between April and June spore concentrations increased up to 9-fold in the first 4–6 wk and then declined. On stem sections of seed plants of oilseed rape and cabbage similarly placed on the soil, the fungi produced viable spores for up to 23 wk with spore concentrations increasing up to 11-fold in the first 6–8 wk after harvest. These results indicate that infected debris of brassica crops remaining on the ground after harvest may provide a source of dark leaf spot infection which may be implicated in the spread of the disease within and between crops.     

RETRACTED ARTICLE: Foraging of Host-Habitat and Superparasitism in <Emphasis Type="Italic">Cotesia glomerata</Emphasis>: A Gregarious Parasitoid of <Emphasis Type="Italic">Pieris brassicae</Emphasis>

The study investigates differences in the oviposition pattern of a braconid parasitoid, Cotesia glomerata (Linn.) in Pieris brassicae (Linn.) in relation to their use of different cruciferous food plants. The response of P. brassicae to superparasitism and consequences for the parasitoid were examined in order to elucidate the ecological significance of this behaviour. Female parasitoid located various crucifers and searched for host more frequently almost on all the host plants tested i.e. cabbage, cauliflower, Chinese cabbage, broccoli and radish. According to the estimated relative number of female locating hosts, cabbage was the most attractive plant for C. glomerata and total number of eggs laid in host larvae feeding on it was higher than in larvae feeding on other plants. Laboratory experiments demonstrated that superparasitism reduced survivorship of P. brassicae larvae. Superparasitism lengthened parasitoid development and prolonged the feeding period of host larvae. Sex ratio and the body weight of emergent wasps correlated negatively with brood size. Despite a trade-off between maximising brood size and optimising the fitness of individual offspring, two or three ovipositions on P. brassicae larvae resulted in a greater female dry mass than did a single oviposition on the host. Thus, superparasitism might be of adaptive significance under certain circumstances, especially when host density is low and unparasitized hosts are rare in a habitat.     

Control of Lepidopteran insect pests in transgenic Chinese cabbage (Brassica rapa ssp. pekinensis) transformed with a synthetic Bacillus thuringiensis cry1C gene

 A synthetic Bacillus thuringiensis cry1C gene was transferred to three Korean cultivars of Chinese cabbage via Agrobacterium tumefaciens-mediated transformation of hypocotyl explants. Hygromycin resistance served as an efficient selective marker. The transformation efficiency ranged from 5% to 9%. Transformation was confirmed by Southern blot analysis, PCR, Northern analysis, and progeny tests. Many transgenic plants of the closed-head types (lines Olympic and Samjin) flowered in vitro. Over 50 hygromycin-resistant plants were successfully transferred to soil. The transgenic plants and their progeny were resistant to diamondback moths (DBM, Plutella xylostella), the major insect pest of crucifers world-wide, as well as to cabbage loopers (Trichoplusia ni) and imported cabbage worms (Pieris rapae). Both susceptible Geneva DBM and a DBM population resistant to Cry1A protein were controlled by the Cry1C-transgenic plants. The efficient and reproducible transformation system described may be useful for the transfer of other agriculturally important genes into Chinese cabbage. Received: 12 June 2000 / Revision received: 21 August 2000 / Accepted: 22 August 2000     

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.     

Plasmodiophora brassicae-induced Cell Death and Medium Alkalization in Clubroot-resistant Cultured Roots of Brassica rapa

Plasmodiophora brassicae causes clubroot in the turnip, Brassica rapa L. We used organ cultures of adventitious roots from B. rapa seedlings to investigate the initial response of resistant and susceptible cultivars to P. brassicae infection. Primary plasmodia of P. brassicae were observed in root hairs of both susceptible and resistant cultured roots. On the other hand, secondary plasmodia were able to proliferate only in the susceptible root culture but not in the resistant one. Root cultures from the susceptible cultivar all developed clubroot 4 weeks after treatment with 10⁴, 10⁵ or 10⁶ spores/ml, but roots from the resistant cultivar did not develop clubroot under the same conditions. Cell death, as measured by Evans blue and TTC dye methods, was observed in cultured roots from the resistant cultivar but did not occur in roots from the susceptible cultivar after exposure to P. brassicae spores. Cell death was inhibited almost completely by EGTA and verapamil but not by the calmodulin antagonist W7. These results suggest the involvement of Ca²⁺ in P. brassicae‐induced cell death. Alkalization of the root culture medium of the resistant cultivar was observed 2 days after treatment with P. brassicae spores but was not observed in root culture medium from the susceptible strain. We conclude that our root culture system must be a useful tool for further studies of the molecular mechanism of clubroot resistance.     

Interspecific hybrids between Brassica napus L. and B. oleracea L. developed by embryo culture

Summary Interspecific hybrids between Brassica napus and B. oleracea are difficult to produce, and previous attempts to transfer economic characters from one species to the other have largely been unsuccessful. In these studies, oilseed rape cv. Tower (2n38) (B. napus) was crossed with broccoli and kale (2n18) (B. oleracea), and hybrid plants were developed from embryos in culture by either organogenesis or somatic embryogenesis. In rape × broccoli, F₁ plants were regenerated from hybrid embryos and the plants produced viable selfed seeds. F₅ plants (2n38) homozygous for white flower colour were selected for high oil content (47%) and Line 15; a selection from these plants produced fertile hybrids with rape, broccoli and kale without embryo culture. In reciprocal crosses between oilseed rape cv. Tower and an aphid resistant diploid kale, 28 and 56 chromosome F₁ hybrid plants were regenerated from somatic embryos. The 56 chromosome plants were self-fertile and it was concluded from F₂ segregation ratios that a single dominant gene controls resistance to cabbage aphid in kale. The 28 chromosome F₁'s were self-sterile, but these and the 56 chromosome F₁'s could be backcrossed to rape and kale. A cross between the F₁ (2n56) and a forage rape resulted in the selection of a cabbage aphid (Brevicoryne brassicae L.) resistant line (Line 3). Both Line 15 and Line 3 can serve as bridges for gene interchange between B. campestris, B. napus and B. oleracea, which has not been possible hitherto. Hybridisations between rape and tetraploid kale produced F₁ plants with 37 chromosomes. One F₂ plant possessed coronal scales and the inheritance was shown to be controlled by a single recessive gene unlinked to petal colour.This paper is dedicated to Mr. T. P. Palmer, a colleague and close friend who retired from the DSIR as Assistant Director of the Crop Research Division in September 1984     

Determining the host‐plant resistance mechanisms for Mamestra brassicae (Lepidoptera: Noctuidae) pest in cabbage

Six cabbage (Brassica oleracea var. capitata) varieties with different levels of resistance to Mamestra brassicae (Lepidoptera: Noctuidae) were investigated in order to assess whether antibiosis and antixenosis mechanisms are involved in the resistance to this pest or not. Four experiments were conducted to determine the effect of variety and plant ontogeny on larval behaviour, adult oviposition and leaf damages in non‐choice and choice tests. Larval survival, time to development and larval weights differed depending on the varieties and plant stages that we tested. At the pre‐head stage, larval mortality was higher, larvae died faster, time to pupation was shorter, pupae were lighter and the percentage of viable pupae and growth index (GI) values were lower than larvae reared from plants at the head stage. The commercial hybrid ‘Corazón de buey’ and the local variety named ‘BRS0535’ exhibited antibiosis to M. brassicae as they reduced its survival and growth and delayed its development time. In addition, these varieties were the most resistant after artificial infestation in terms of head foliage consumption and number of larvae per plant. Oviposition tests demonstrated that resistance found in ‘Corazón de buey’ and BRS0535 could be also based on antixenosis mechanisms as they resulted in fewer egg batches on plants, whereas BRS0402 could be classified as resistant because M. brassicae larvae showed less preference for it. Thus, resistance to M. brassicae found in cabbage crops may be due to the joint action of several factors involving antibiosis and antixenosis. We found significant differences in the resistance of BRS0535 depending on the plant ontogeny as it loses its resistance while developing. Further studies are required to identify the mechanism of antibiotic resistance which is present in this variety at the pre‐head stage and the changes that occur in plant defence as it grows.     

A behavioural study to help clarify how undersowing with clover affects host-plant selection by pest insects of brassica crops

Laboratory and field-cage tests were done to determine how undersowing brassica plants (Brassica oleraceae L. and B. rapa L.) (Cruciferae) with subterranean clover (Trifolium subterraneum L.) (Papilionaceae) affected host-plant selection by eight pest insect species of brassica crops. The pest species tested were Pieris rapae (Lepidoptera: Pieridae) (the small white butterfly), Pieris brassicae (Lepidoptera: Pieridae) (the large white butterfly), Delia radicum (Diptera: Anthomyiidae) (the cabbage root fly), Phaedon cochleariae (Coleoptera: Chrysomelidae) (the mustard beetle), Plutella xylostella (Lepidoptera: Yponomeutidae) (the diamond-back moth), Evergestis forficalis (Lepidoptera: Pyralidae) (the garden-pebble moth), Mamestra brassicae (Lepidoptera: Noctuidae) (the cabbage moth) and Brevicoryne brassicae (Hemiptera: Aphididae) (the cabbage aphid). In all tests, except two in which the brassica plants were about three times as high as the clover background, 39%–100% fewer of the pest insect stage monitored were found on host plants presented in clover than on those presented in bare soil. Contrary to claims supporting the ‘enemies hypothesis’, differences in colonization alone appeared sufficient to account for the lower numbers of insects found when host plants are undersown with clover. To be effective in reducing plant colonization, the clover must cover 50%, and preferably more, of the vertical profile of the crop plants. As clover used as an undersown crop often has to be cut to make it less competitive with the main brassica crop, temporal aspects of the condition of the clover during critical periods of pest activity need to be recorded carefully before concluding that undersowing does not produce the effect desired against certain pest species under field conditions. The effective clover barrier is like any other treatment, if it is not present at the appropriate time it cannot be expected to reduce pest insect numbers.     

Falling aphids enhance impact of biological control by parasitoids on partially aphid-resistant plant varieties

Cage experiments, in which the population increase of Metopolophium dirhodum was measured either in the absence or presence of the parasitoid Aphidius rhopalosiphi, showed reductions of peak populations in the presence of the parasitoid of 30% on the susceptible wheat cultivar Armada, but of 57% on the partially resistant Rapier. On both cultivars, aphid population curves in the presence and absence of the parasitoid diverged before the first mummies were formed. The number of aphids leaving plants was measured, and was found to increase in the presence of the parasitoid, particularly with Rapier (almost double). Most aphids (about 75%) which left plants fell, and a smaller proportion regained plants when parasitoids were present than when they were absent. A field experiment with Brevicoryne brassicae and natural predation on Brussels sprouts also showed an increase in aphids leaving a less susceptible compared with a more susceptible cultivar.     

Cytology of infection,development and expression of resistance to Plasmodiophora brassicae in canola

The timing and expression of resistance to four isolates of Plasmodiophora brassicae, collected from research sites where pathotypes 2, 3, 5 and 6 (Williams' system) had been dominant when characterised in 2006, were assessed in four new commercial cultivars of canola (Brassica napus) with resistance to clubroot. Each of the resistant cultivars was highly resistant to all four of the isolates, and there was no difference in their response to infection. Root hair infection occurred at high levels, but pathogen development occurred more slowly than in a susceptible cultivar (control). Secondary infection and development in cortical cells was severely inhibited in each of the resistant cultivars; only a few bi‐nucleated plasmodia were observed at 12 days after inoculation (DAI), and plasmodia were rarely observed at 18 and 24 DAI. In contrast, development in the susceptible cultivar had progressed to resting spores by 24 DAI. A dense ring of accumulated reactive oxygen species (ROS) was observed in the endodermis, pericycle and vascular cambium of non‐inoculated controls and inoculated plants of the resistant cultivars. However, the ROS ring disappeared rapidly in infected plants of the susceptible cultivar. Plasmodia invaded the stele of susceptible roots by preferentially colonising the xylem parenchyma cells. Expansion and enlargement of lignified xylem cells was observed by 35 DAI. The absence of any specific points of ROS accumulation or lignification of epidermal or cortical cells in the resistant cultivars indicates that a hypersensitive response is not the main mechanism of resistance in these lines. The uniform response of these resistant cultivars to the four isolates of P. brassicae indicates that the resistance in each cultivar may be conditioned by a gene(s) from a single source that confers broad resistance, because most sources of resistance to P. brassicae are pathotype specific.     

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.