Revisiting strategies for reducing the seedbank of Orobanche and Phelipanche spp.

Weedy Orobanche and Phelipanche species are important in Southern and Eastern Europe, the Middle East and North Africa, and have recently been reported in the USA, Australia and some countries in South America. Rather than being controlled, the Orobanche and Phelipanche problem is increasing, both in intensity and in acreage. Large areas of new territory are at risk of invasion, if care is not immediately taken to limit the introduction of parasitic seeds and to educate farmers and others to be alert for new infestations. There is an urgent need to re-evaluate novel integrated Orobanche and Phelipanche management programmes that will allow a better control of the parasite species and limit their distribution. The main obstacle in the long-term management of Orobanche and Phelipanche infested fields is the durable seedbank, which may remain viable for decades in the field. Large quantities of long-lived seeds give the parasite genetic adaptability to changes in host resistance and cultural practices. As long as the seedbank is not controlled, the need to control the parasite will persist whenever a susceptible host is grown in the infested field. We discuss strategies for seedbank reduction, paying particular attention to cultural practices, whereas chemical and biological control methods are covered by other reviews in this issue.     

Biology and management of the fast‐emerging threat of broomrape in rapeseed–mustard

Broomrapes (Phelipanche and Orobanche spp.) are obligate root parasites of the family Orobanchaceae. The natural variation in Orobanchaceae exists in plants involving Triphysaria versicolor (facultative hemiparasite), Striga hermonthica (obligate hemiparasite) and Phelipanche aegyptiaca, formerly Orobanche aegyptiaca (holoparasite). The family Orobanchaceae has 90 genera. The four major parasitic species of broomrape are Phelipanche ramosa in the Brassicas, especially in rapeseed, Phelipanche cernua in tobacco and sunflower, P. aegyptiaca in solanacious crops and in Brassica juncea and Phelipanche crenata in leguminous crops. They are prevalent, infesting nearly 1.2% of the world's arable land. In India, P. ramosa and P. aegyptiaca cause severe infestations of Brassica and have threatened its cultivation in major growing areas. In addition, a single broomrape plant can release >100 000 seeds that remain viable for decades in the soil. This provides the parasite with a great genetic adaptability to environmental changes, including host resistance, agronomical practices and herbicide treatments. Different control measures, such as manual weeding, herbicide applications, solarization, crop rotation and integrated broomrape management practices, have been proposed in order to reduce the broomrape menace and improve yields in farmers' fields. Breeding for broomrape resistance also could be an economic, feasible and environmental friendly control method. The present article reviews the current status of research on broomrape in India and abroad, as well as suggests strategies for its effective management.     

Strigolactone deficiency confers resistance in tomato line SL-ORT1 to the parasitic weeds Phelipanche and Orobanche spp

The parasitic flowering plants of the genera Orobanche and Phelipanche (broomrape species) are obligatory chlorophyll-lacking root-parasitic weeds that infect dicotyledonous plants and cause heavy economic losses in a wide variety of plant species in warm-temperate and subtropical regions. One of the most effective strategies for broomrape control is crop breeding for broomrape resistance. Previous efforts to find natural broomrape-resistant tomato (Solanum lycopersicon) genotypes were unsuccessful, and no broomrape resistance was found in any wild tomato species. Recently, however, the fast-neutron-mutagenized tomato mutant SL-ORT1 was found to be highly resistant to various Phelipanche and Orobanche spp. Nevertheless, SL-ORT1 plants were parasitized by Phelipanche aegyptiaca if grown in pots together with the susceptible tomato cv. M-82. In the present study, no toxic activity or inhibition of Phelipanche seed germination could be detected in the SL-ORT1 root extracts. SL-ORT1 roots did not induce Phelipanche seed germination in pots but they were parasitized, at the same level as M-82, after application of the synthetic germination stimulant GR24 to the rhizosphere. Whereas liquid chromatography coupled to tandem mass spectrometry analysis of root exudates of M-82 revealed the presence of the strigolactones orobanchol, solanacol, and didehydro-orobanchol isomer, these compounds were not found in the exudates of SL-ORT1. It can be concluded that SL-ORT1 resistance results from its inability to produce and secrete natural germination stimulants to the rhizosphere.     

In vitro culture of Orobanche ramosa

Orobanche spp. (broomrapes) are holoparasites that subsist on the roots of many important crops and can considerably reduce yield. The control of Orobanche spp. includes physical, chemical and biological methods. Interactions between parasitic angiosperms and their hosts first occur at the level of parasite seed germination. The seeds of all Orobanchaceae germinate in soil under natural conditions only in response to specific chemical exudates from the host plant. This study describes the influence of different plant growth regulators and host plant root exudates on germination and development of calli from Orobanche seeds in vitro . The effect of indole-3-acetic acid, gibberellic acid and kinetin on the germination of Orobanche seeds varied with concentration. These plant growth regulators also affected the period of germination and the structure of calli and protrusions. An in vitro system for the collection of tobacco root exudates was established. Compounds released from the host roots of three different tobacco cultivars were found to provoke high levels of germination of the Orobanche seeds without any period of pre-conditioning. This study developed methods for the investigation of host–parasite interactions and the effect of germination stimulants in Orobanche spp.     

新疆列当种类、危害与防治现状

列当是一种根部全寄生恶性杂草,影响作物生长,对农业生产造成了巨大破坏。目前,列当防治措施主要包括轮作、种植诱捕作物、化学防治等,但尚无适用于大部分地区及作物的有效防治措施。本文综述了新疆列当种类、寄生特点、危害和目前采取的防治措施,并对列当的萌发刺激物、防治机理和化学除草剂在大田应用提出展望,以期为新疆相关防治工作提供参考。     

Ausbreitung des parasitischen Unkrauts Phelipanche ramosa in der deutschen Landwirtschaft

The root parasitic weeds Phelipanche ramosa (branched broomrape) and P. aegyptiaca have the widest host range among Orobanche and Phelipanche species. In Western Europe, P. ramosa attacks, with increasing aggressiveness, crops such as oilseed rape, tobacco, hemp, and tomato. The unique biology of root parasites, establishes a closed link with their host plant, thus reducing the possibility to successfully control them. Control measures include (a) physical processes (such as weeding, solarization, deep ploughing, burning off, flooding), (b) chemical (like soil fumigation, use of herbicides, germination stimulants) and (c) biological methods (e.g. resistant varieties, use of fungi and insects as antagonists, trap and catch crops). German tobacco growers rely mostly on the herbicide method. They apply glyphosate in very low concentrations, when the first tubercles are formed. Also a fungal antagonist against the parasitic weed on tobacco was found in Germany, but until now has not been developed into a commercial mycoherbicide. After hemp production lost its significance as a crop in Germany, tobacco remained as the main host for P. ramosa. In the past 10 years, branched broomrape has spread out and currently it can be found in areas where previously were free of it. Since the elimination of EU subsidies, some tobacco growers began to cultivate on their land parsley instead of tobacco. As a result, parsley has now been infected with P. ramosa. When used 10 years ago as catch crop, parsley had a rather small effect on branched broomrape. This potential danger, especially by other potential host plants, such as oilseed rape, tomato and potato or even weeds should not be underestimated. Spread and expansion of the host plant spectrum of branched broomrape in Germany might be reduced by the introduction of appropriate phytosanitary measures and improved information policies.     

列当(Orobanche spp. and Phelipanche spp.)种子的采集与预处理方法

列当(Orobanche spp.and Phelipanche spp.)是一种在世界范围内危害严重的根寄生杂草,防除列当的研究也越来越多。正确地采集和预培养列当种子是获得正确防除列当研究结果的前提。列出了大田正确采集列当种子的方法,以及对采集到的种子进行过筛提纯、加洗洁精和吐温20清洗后,再表面消毒和预培养,并通过发芽试验检验此方法的可行性。结果表明,经过处理的向日葵列当、瓜列当种子的发芽率最高分别为54.4%、59.1%。此方法可以应用于列当的采集和预处理过程中,可为列当研究和防除提供参考。     

Factors affecting the efficacy of Orobanche cumana chemical control in sunflower

Broomrapes ( Orobanche spp.) are root parasites, causing severe damage to vegetable and field crops. Their early in situ detection for efficient control is crucial but difficult, because of their sub-surface development. Minirhizotron technology enables the in situ observation of host–parasite interactions. The initial underground development of Orobanche cumana on sunflower roots and its control with imazapic were investigated. Imazapic (2.4 g active ingredient ha⁻¹) was applied post-emergence on sunflower at the eight-true-leaf stage, when O. cumana attachments were first observed. No viable or new attachments were detected on roots of the treated sunflower after herbicide application. However, within the same time frame, the already existing O. cumana attachments in the non-treated controls continued to develop and new attachments appeared. Orobanche cumana shoot emergence was correlated with growing day-degrees (GDD, d °C). This correlation was quantified by a four-parameter logistic equation. Control efficacy was affected by depth and number of parasite attachments per minirhizotron camera frame: it was highest in the topmost layer, and decreased with depth and number of attachments. The results obtained in this study emphasise the need to develop a decision support system based on GDD that should be validated under long-term field conditions.     

列当杂草及其防除措施展望

根寄生杂草列当Orobanche spp.每株可产生大量细小的种子,易于传播,且在出苗前已经对寄主作物造成危害,使列当的防除成为一个世界性难题。本文详述了列当的生活史、为害现状以及防除措施,并突出了生物防除的生态效益与经济效益。基于＂诱捕＂作物及＂捕获＂作物诱导列当种子＂自杀发芽＂理论的防除机制,本文提出了新型可持续生态防除思想,为列当科杂草防除提供参考。     

Study of Resistance to Orobanche ramosa in Host (Oilseed Rape and Carrot) and Non-host (Maize) Plants

Orobanche ramosa is a parasitic Angiosperm responsible for severe yield losses in several economical crops. It is a serious threat in oilseed rape in France and Morocco and is appearing in carrot crops in Morocco. In this study, several varieties of oilseed rape and carrot were screened in order to identify resistant cultivars and to characterize the resistance mechanisms involved. All the 15 oilseed rape varieties tested were susceptible. In carrot, the varieties 'Colmar à coeur rouge' and 'Nantaise demi-longue' were susceptible, whereas 'Palaiseau' and 'Buror' were resistant. In the susceptible 'Colmar à coeur rouge' carrot no defence reactions were found and the development of the parasite inhibited carrot tap root formation. In the resistant carrot varieties, the parasite germinated, became attached to the host root but became necrotic before emergence. In 'Buror' carrot, formation of a mechanical barrier was associated with the restriction to the cortex of the parasite. In maize cv. 'Vigni', a non-host of O. ramosa, thickening of xylem vessels, cell divisions in the central cylinder and formation of an encapsulation layer were observed in association with restricted development of Orobanche haustoria.     

Comparative proteomic analysis of Orobanche and Phelipanche species inferred from seed proteins

Orobanche and Phelipanche species are root parasitic plants some of which infect important crops, causing extensive damage. In an attempt to unravel molecular systematic in these genera using protein markers, we established the protein profile of twelve species by two-dimensional gel electrophoresis (2-DE). Proteins were detected by Sypro Ruby staining and the gel images were digitalised and analysed. The 2-DE maps contained from 30 to 140 protein spots for the analysed species. A virtual master gel was built with spots present in all species and revealing the presence of 260 protein spots. Only one protein was monomorphic, while 143 were unique to a single species. Proteins were scored for the presence or absence of homologous spots to create a binary matrix. Phylogenetic analysis was carried out using maximum parsimony and distance method. The trees obtained were rooted using Cistanche phelypaea as an out-group. Both analyses showed Orobanche as a monophyletic group. AMOVA analysis revealed phenotypic diversity between species within genera, as well as a considerable level of variation between genera. This is a first step to help in the phylogenetic classification of parasitic plants by means of proteomics.     

Understanding Orobanche and Phelipanche–host plant interactions and developing resistance

Orobanche and Phelipanche spp. (broomrapes) are parasitic plants that can be responsible for devastating losses in several important crops. The development of resistant cultivars is one of the key strategies in the fight against this pest. However, the nature of resistance is complex and the basis of the interaction between the host and the parasite is still largely unknown. Despite the progress achieved during the last century through breeding programmes, sources of resistance are often scarce (e.g. the legumes). The resistance that is available is often not durable, with field resistance being overcome by new races of the parasite (e.g. sunflower). This review summarises efforts made to improve the resistance of crop hosts for broomrapes through classical breeding programmes and looks forward to the integration of new knowledge generated from molecular and morphological studies. Emphasis is given to the need for a multidisciplinary approach to achieve success, ranging from improved field phenotyping to genetic and biotechnological studies. All components are necessary to understand this particular and characteristic interaction: a plant parasitising another plant.     

Use of zoophytophagous mirid bugs in horticultural crops: Current challenges and future perspectives

In recent years, the use of predatory mirid bugs (Hemiptera: Miridae) in horticultural crops has increased considerably. Mirid bugs are zoophytophagous predators, that is, they display omnivorous behavior and feed on both plants and arthropods. Mirid bugs feed effectively on a wide range of prey, such as whiteflies, lepidopteran eggs and mites. In addition, the phytophagous behavior of mirid bugs can activate defenses in the plants on which they feed. Despite the positive biological attributes, their use still presents some constraints. Their establishment and retention on the crop is not always easy and economic plant damage can be caused by some mirid species. In this review, the current strategies for using zoophytophagous mirid bugs in horticultural crops, mainly Nesidiocoris tenuis, Macrolophus pygmaeus and Dicyphus hesperus, are reviewed. We discuss six different approaches which, in our opinion, can optimize the efficacy of mirids as biocontrol agents and help expand their use into more areas worldwide. In this review we (i) highlight the large number of species and biotypes which are yet to be described and explore their applicability, (ii) present how it is possible to take advantage of the mirid‐induced plant defenses to improve pest management, (iii) argue that genetic selection of improved mirid strains is feasible, (iv) explore the use of companion plants and the use of alternative foods to improve the mirid bug management, and finally (vi) discuss strategies for the expansion of mirid bugs as biological control agents to horticultural crops other than just tomatoes. © 2020 Society of Chemical Industry     

Seed viability determination in parasitic broomrapes (Orobanche and Phelipanche) using fluorescein diacetate staining

Fluorescein diacetate (FDA) was investigated as a potential viability stain for seeds of holoparasitic broomrapes Orobanche and Phelipanche (Orobanchaceae), using four weedy and two non-weedy taxa. FDA viabilities were compared with the currently used 2,3,5-triphenyl tetrazolium chloride (TTC) test, and in vitro germination induced by the strigolactone analogue GR-24. Viable FDA-stained seeds were characterised by bright yellow–green fluorescence under 450–490 nm blue light. These viable seeds could be discriminated from non-viable seeds, in which low levels of fluorescence, similar to non-stained control seeds were observed. FDA viabilities were not significantly different from those obtained using the TTC stain; however, viability values for both stains were consistently higher than levels of artificially induced germination. Positive TTC-staining showed continuous variation that was difficult to interpret. Nevertheless, the TTC test does not involve grinding seeds, and therefore probably remains a practical alternative to FDA for screening soil samples contaminated with Orobanche and Phelipanche seeds, which indicates different applications for the two viability tests. Interestingly, the strigolactone analogue GR-24 only induced germination in O. crenata , P. ramosa and O. minor , suggesting a high specificity towards this germination inducer among the species investigated. We suggest that FDA-staining provides a potential alternative to the currently used TTC test for seed viability assays for Orobanche and Phelipanche , and other parasitic weeds which are an obstacle to crop cultivation.     

储藏物害虫生物性防治技术研究现状和展望

有效的杀虫剂和熏蒸剂种类的减少,以及它们对环境的污染和对人畜健康的影响,使得生物性的防治技术在储藏物害虫的控制上显得越来越重要。近年来,有关储藏物害虫生物性防治技术的研究报道越来越多。根据过去20年的文献,特别是根据刚出版的第七届国际储藏物保护会议的有关论文,评述了储藏物害虫生物性防治技术的研究现状,展望了将来的发展。内容涉及到天敌昆虫、昆虫信息素、病原微生物、抗虫品种和遗传防治等领域,着重于它们在实践中的应用。     

An Integrated Approach in the Control of Potato Root Eelworm

During the last decade, new control possibilities for the potato root eelworm have come to the fore, namely resistant potato varieties and soil disinfection. These have led to an integrated approach to the control of the potato root eelworm in the Netherlands. For this purpose, a number of potato rotations, aiming at the prevention or elimination of non-detectable populations of the potato root eelworm, are prescribed for non-infested fields. These regulations allow the farmer to grow potatoes more frequently on non-infested soils if he uses control measures (i.e. resistant potato varieties, soil disinfection). More stringent restrictions are in force for infested fields on which the growing of susceptible crops and propagation material is, of course, prohibited.     

Fenugreek root exudates show species-specific stimulation of Orobanche seed germination

Various Orobanche species are weedy and cause severe reduction in the yields of many important crops. The seeds of these parasitic weeds may remain dormant in the soil for many years until germination is stimulated by the release of a chemical signal from a host plant. In order to determine the effects of fenugreek root exudate on the induction of Orobanche crenata, Orobanche ramosa and Orobanche foetida seed germination, root exudate was collected from hydroponically grown fenugreek seedlings. Fractionation patterns obtained from column and thin layer chromatography of the fenugreek root exudate showed a set of metabolites differing in their polarity with stimulatory activity on Orobanche seed germination. The crude root exudate stimulated both O. ramosa and O. crenata seed germination to the same level caused by the synthetic germination stimulant GR24 at 10 mg L^?1. It also stimulated O. foetida seed germination which did not respond to GR24. Active fractions of root exudate stimulated the germination of Orobanche species differentially.     

Use of ITS nuclear sequences from Phelipanche aegyptiaca as a direct tool to detect single seeds of broomrape species in the soil

Broomrape (Phelipanche and Orobanche spp.) are obligate holoparasites that attack roots of almost all economically-important crops in semiarid regions of the world. Broomrape seeds are extremely small (dust-like seeds), averaging 200 to 300?μm in size and because of the miniscule seed size it is difficult to detect and confirm via conventional methods. In this study our aim was to develop a PCR-based assay specific for broomrape soil-borne seeds and sensitive enough to detect a single or few broomrape seeds in a soil sample. For this purpose, we used complementary polymerase chain reaction (PCR) primers based upon unique sequences in the internal transcribed spacer (ITS) regions of the nuclear ribosomal DNA of Phelipanche aegyptiaca. Genomic DNA was extracted from soil samples artificially infested with broomrape seeds or tissue of Phelipanche aegyptiaca Pers., Orobanche cumana Wallr. and Phelipanche crenata Forsk. and subjected to PCR analysis. Using ITS-350 primers, a specific PCR product (350?bp) was amplified and detected in all samples containing broomrape species, but was not detected in soil sample free of broomrape seeds or tissues. Additionally, the PCR-based assay was sensitive enough to detect even a single broomrape seed in the soil. As expected the universal internal control primers amplified a PCR product (555?bp) of genomic DNA extracted from soil samples with or without broomrape tissues or seeds. This diagnostic method is simple, reliable and rapid and could help for assessment of broomrape seed contamination in a crop field.     

Potato leafroll virus (PLRV): its transmission and control

This review provides a wide assessment of the present state of knowledge about the potato leafroll luteovirus (PLRV), a pathogen which is seriously devastating potato crops in many parts of the world. The main biological, physical and chemical properties of this virus are described. The transmission of PLRV by aphids, which are the only transmitters of this virus under natural conditions, is characterized. Special attention is given to the control of PLRV through the use of resistant potato cultivars. Recent advances in obtaining resistant transgenic plants are outlined.     

Pleiotropic effects of herbicide‐resistance genes on crop yield: a review

The rapid adoption of genetically engineered herbicide‐resistant crop varieties (HRCVs)—encompassing 83% of all GM crops and nearly 8% of the worldwide arable area—is due to technical efficiency and higher returns. Other herbicide‐resistant varieties obtained from genetic resources and mutagenesis have also been successfully released. Although the benefit for weed control is the main criteria for choosing HRCVs, the pleiotropic costs of genes endowing resistance have rarely been investigated in crops. Here the available data of comparisons between isogenic resistant and susceptible varieties are reviewed. Pleiotropic harmful effects on yield are reported in half of the cases, mostly with resistance mechanisms that originate from genetic resources and mutagenesis (atrazine in oilseed rape and millet, trifluralin in millet, imazamox in cotton) rather than genetic engineering (chlorsulfuron and glufosinate in some oilseed rape varieties, glyphosate in soybean). No effect was found for sethoxydim and bromoxynil resistance. Variable minor effects were found for imazamox, chlorsulfuron, glufosinate and glyphosate resistance. The importance of the breeding plan and the genetic background on the emergence of these effects is pointed out. Breeders' efforts to produce better varieties could compensate for the yield loss, which eliminates any possibility of formulating generic conclusions on pleiotropic effects that can be applied to all resistant crops. © 2013 Society of Chemical Industry