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.     

Phelipanche aegyptiaca management in tomato

Phelipanche and Orobanche species (broomrapes) are root holoparasitic plants that cause severe damage to economically important crops. Phelipanche and Orobanche spp. are widespread in Mediterranean areas, in Asia and in Southern and Eastern Europe, attacking dicotyledonous crops and depending entirely on their hosts for all nutritional requirements. Phelipanche aegyptiaca , Phelipanche ramosa and Orobanche cernua are extremely troublesome weeds on tomatoes. These noxious parasites exert their greatest damage prior to their shoot emergence and flowering; therefore, the majority of field loss may occur before diagnosis of infection. This review summarises the four main control measures for the weedy root parasites Phelipanche and Orobanche in tomato, namely chemical and biological control, resistant varieties and sanitation. Some of these methods are commercially widely used by farmers in Israel (chemical control), some are in the final stages of development towards commercialisation (resistant varieties and sanitation), and some still require further development and improvement before commercial implementation (biological control). The review presents an up-to-date summary of the available knowledge on their use for broomrape management in processing tomatoes.     

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。     

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.     

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.     

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。     

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.     

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.     

Parasitic weeds of the Orobanchaceae family and their natural hosts in Jordan

A field survey of the Orobanchaceae family members and their hosts in Jordan was carried out from 2003 to 2007. The intensity of parasite infection on different hosts and the severity of the infestation were evaluated. The results showed the presence of seven species of Orobanche and three species of Cistanche . The Orobanche species were found parasitizing 86 plant species belonging to 24 botanical families. Most of the species attacked by Orobanche were from the Compositae (20 species), Solanaceae (11 species), Leguminosae (nine species), Umbelliferae (seven species), Cruciferae (seven species), Cucurbitaceae (four species), Labiatae (four species), and Rosaceae (four species) families. Other families were represented by one-to-three species. Cistanche attacked 20 species of forage wild shrubs, fruit trees, and forest trees of seven families, mostly belonging to the Chenopodiaceae (seven species) and Leguminosae (three species) families. Previously unreported hosts for both genera include: Amygdalus communis , Olea europaea , and Quercus coccifera , which were parasitized by Orobanche palaestina ; A. communis , O. europaea , Prunus armeniaca , and Prunus persica , which were parasitzed by Orobanche cernua ; O. europaea and A. communis , which were parasitzed by Orobanche schultzii ; Haloxylon persicum , which was parasitzed by Cistanche lutea ; Punica granatum , Alhagi maurorum , Casuarina equisetifolia , Centaurea postii , and Prosopis farcta , which were parasitzed by Cistanche tubulosa ; and Achillea spp., Anabasis syriaca , H. persicum , Haloxylon salicornicum, Suaeda spp., and Zilla spinosa , which were parasitzed by Cistanche salsa . Certain Orobanche species were completely destructive to the cultivated crops. The results indicated the high potential of both parasitic genera to spread and to attack new hosts, while the threat they impose to agriculture in Jordan will probably result from poor management and deficiences in farmers' training.     

Characterization of the Orobanche–Medicago truncatula association for studying early stages of the parasite–host interaction

There is an increasing interest in the legume species Medicago truncatula as a model plant for structural and functional genomic studies that can be used to identify agronomically important genes in legumes. Field screening has shown very high levels of resistance to Orobanche crenata in M. truncatula. However, in vitro studies with O. crenata, Orobanche foetida, Orobanche ramosa and Orobanche minor showed useful variation among accessions at early stages of the parasite–host interaction. Significant differences were observed in the levels of germination of O. crenata and O. foetida seeds induced by different accessions of M. truncatula. Only limited germination was observed on accession SA‐4327. All accessions induced little O. ramosa and O. minor germination. Accessions also varied in the number of O. crenata and O. foetida attachments supported, with few developing on accession SA‐27774. The variation observed for induction of germination and of subsequent attachment will be useful to isolating and characterizing genes involved in the early stages of Orobanche–host plant interaction and for the study of the biosynthetic pathways of production for germination stimulants.     

Quantitative aspects of Orobanche crenata infestation in faba beans as affected by abiotic factors and parasite soil seedbank

The interactions between the root parasitic weed Orobanche crenata Forsk. and its host plant faba bean ( Vicia faba L.) were quantified under controlled and field conditions at ICARDA's Tel Hadya research station. In the field experiments conducted in 1993–94 and 1994–95 faba beans were sown on two dates, in plots with 0, 50, 200 and 600 O. crenata seeds kg^–1 soil, under both limited and sufficient moisture supply. The effects of temperature on the duration of the early developmental stages of O. crenata were investigated in a growth chamber. The extent of O. crenata infestation was closely related to the number of parasite seeds in the soil. The seed-density treatment with 600 seeds kg^–1 soil resulted in complete crop failure. Furthermore, O. crenata infestation was higher under sufficient than under limiting water supply conditions, irrespective of sowing date. Only in the moderately infested plots, did shifting of the planting time of faba bean result in a significant decrease in parasite dry weight and an increase in crop seed yield. The timing of germination, attachment and further developmental stages of O. crenata was not related to faba bean growth stage and was affected primarily by soil temperature. The duration of O. crenata developmental stages was estimated using the thermal time concept. The relationship between total number of parasite attachments at the harvest of the faba bean crop and O. crenata seed density was dependent on maximum faba bean root-length density measured by the start of pod-filling in each treatment combination of sowing date and moisture supply. The results are discussed with reference to implications for the development of a dynamic simulation model for the prediction of faba bean yield losses caused by O. crenata .     

Focus on ecological weed management: what is hindering adoption?

Despite increased concerns regarding the heavy reliance of many cropping systems on chemical weed control, adoption of ecological weed management practices is only steadily progressing. For this reason, this paper reflects on both the possibilities and limitations of cultural weed control practices. Cultural weed control utilises a number of principles, predominantly: (i) a reduced recruitment of weed seedlings from the soil seedbank, (ii) an alteration of crop–weed competitive relations to the benefit of the crop and (iii) a gradual reduction of the size of the weed seedbank. Compared with chemical control, the general applicability, reliability and efficacy of most measures is only moderate, and consequently, cultural control strategies need to consist of a combination of measures, resulting in increased systems complexity. Combined with the trade‐offs connected to some of the measures, this hampers large‐scale implementation. It is argued that tailoring cultural weed management strategies to the needs and skills of individual farmers would be an important step forward. Research can aid in improving the utilisation of cultural weed control strategies by focussing on a broadening of the range of available measures and by providing clear quantitative insight in efficacy, variability in outcome and trade‐offs of these measures.     

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.     

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。     

Purple witchweed (Striga hermonthica) germination and seedbank depletion under different crops,fallow, and bare soil

Seedbank density is an important aspect that determines the amount of damage that the parasitic weed, purple witchweed (Striga hermonthica; hereafter, called “Striga”), causes on its crop hosts. The seedbank depletion of Striga was measured in Mali and Niger during the 2004 rainy season under the host crops, pearl millet and sorghum, the non‐host crops, cowpea and sesame, the intercrops of pearl millet or sorghum with cowpea or sesame, and fallow with or without weeding. Two methods were used and compared; namely, a seed bag method and a soil‐sampling method. The fate of the seeds was assessed by a seed press test. Seed germination, as determined by the presence of empty seed coats, contributed most to the seedbank depletion of Striga under a variety of crop covers and fallow. The highest seedbank depletion was found under the monocultures of the host crops. The intercrops of the host and non‐host crops caused less seedbank depletion, followed by the monocultures of the non‐host crops, fallow, and bare soil. The seed bag method and the soil‐sampling method yielded similar percentages of seedbank depletion, while the former allowed for distinguishing between the germinated and diseased seeds. The results suggest that, although all the tested crop species can cause the seed germination and seedbank depletion of Striga, management by using host cereal crops causes the highest amount of germination and has the highest potential to deplete the soil seed bank, provided that seed production is prevented.     

Investigation of wheat as a trap crop for control of Orobanche minor

Orobanche minor is a parasitic weed that attaches to the roots of red clover (Trifolium pratense) and a number of other broad‐leaved plant species in the Pacific Northwest USA. Orobanche minor seed must be stimulated by host plant exudates for germination and attachment to occur. However, plant species called false‐hosts can stimulate parasitic seed germination without attachment. These species could be utilized as trap crops to reduce the amount of parasitic seed in infested soil. Wheat (Triticum aestivum), was found to be a false‐host of O. minor; therefore, growth chamber, glasshouse and field soil experiments were conducted to evaluate the effect of six soft white winter wheats (T. aestivum), one durum wheat (Triticum turgidum), and one triticale (Triticale hexaploide) on O. minor germination. In growth chamber experiments, wheat and triticale induced 20–70% of O. minor seeds to germinate. In glasshouse studies, O. minor attachment was minimal on red clover plants grown in pots previously planted to wheat or triticale. In pots that did not receive a false‐host treatment, red clover plants averaged 4.2 O. minor attachments per plant. Red clover plants also had fewer O. minor attachments when grown in field soil taken from the plots where wheat or triticale were grown compared with plants grown in soil where no wheat or triticale were previously grown. Our results demonstrate that wheat may have the potential to be effectively integrated into an O. minor management system.     

Dependence of weed flora on the active soil seedbank

Previous studies have shown that reliable predictions of the above-ground weed flora may not be obtainable by using the total number of seeds in the seedbank. The purpose of this study was to determine if the actual weed flora depends on the active soil seedbank (seeds that are germinable in the spring). In the 1995 growing season, seedling emergence was monitored within permanent quadrats established in a field. Soil cores taken from the same field were also monitored for seedling emergence in a greenhouse to estimate the number of seeds in the active soil seedbank. Significant positive relationships were observed between the above two variables based on either total weed flora or individual weed species commonly found in south-western Ontario, Canada. For relatively large-seeded species, the relationship held with a sampling depth up to 15 cm below the soil surface. For small-seeded species, a shallow sampling depth (0–7·5 cm) generated better results than deeper sampling (0–15 cm). In general, 3–7% of the seeds in the active soil seedbank were capable of producing seedlings in the field. The results suggest that the level of weed infestation in a growing season may be predicted using seeds in the active soil seedbank.     

Minimal soil disturbance combined with spring cropping can halt soil seedbank accumulation of Alopecurus myosuroides

The basic mechanism of soil inversion tillage for control of annual weeds is based on the vertical translocation of weed seeds from the soil surface to deeper soil layers. Buried weed seeds either remain dormant in the soil seedbank and are exposed to biological and chemical decay mechanisms, or they germinate but the seedlings cannot reach the soil surface (fatal germination). However, depending on the seed biology of the respective target species, frequent inversion tillage can lead to a build-up of the soil seedbank. For soil seedbank depletion based on available knowledge of the biology of Alopecurus myosuroides seeds, soil inversion tillage is suggested to be reduced to every third or fourth year with reduced or even no-tillage (direct seeding) in between (rotational inversion tillage systems). Including spring crops in the crop rotation could further help dampening the population growth and hence the seed return into the seedbank. This study investigated the effect of rotational inversion tillage in combination with reduced tillage or direct seeding on the soil seedbank and population development of A. myosuroides. In a long-term field trial, set up in 2012, these tillage strategies were compared with continuous inversion tillage in a 3-year crop rotation with two consecutive years of winter wheat (Triticum aestivum) followed by spring barley (Hordeum vulgare). The results showed a significant decline in the soil seedbank following the spring crop, irrespective of the tillage system. The continuous inversion tillage system and inversion tillage before spring cropping with reduced tillage (shallow tillage with a disc harrow) before winter wheat both led to accumulation of seeds in the soil seedbank. In contrast, inversion tillage before spring cropping with direct seeding of winter wheat depleted the soil seedbank significantly after only one crop rotation. Although only covering one intensively studied field site, these findings highlight the need for diversified cropping systems and indicate potential avenues for reducing soil tillage while controlling economically important weeds.     

Weed management strategies for the control of Rottboellia cochinchinensis in maize in Trinidad

Summary. A weed management strategy for the control of Rottboellia cochinchinensis (Lour.) W.D. Clayton in maize, based on reducing the level of spikelet germination from the seedbank and reducing the intensity of weed-crop interference, was conducted over two seasons. The management practices evaluated included chemical control (atrazine, metolachlor, pendimethalin, EPTC), cultural control (straw mulch), mechanical control (inter-row cultivation) and biological control (living mulch). The results indicated that the various management practices had no effect on the growth and development of the crop but significantly affected the weed. The pendimethalin (1°5 kg a.i. ha⁻¹) and inter-row cultivation (14 and 28 days after planting) treatments gave satisfactory control of the weed in the crop over the duration of the critical period of weed competition. All other treatments were ineffective. All management practices made significant contributions to the weed seedbank, thus ensuring the predominance of the weed.     

Effects of host plant genotype and seedbank density on Striga reproduction

Prevention of seed input to the seedbank of Striga hermonthica‐infested fields is an important objective of Striga management. In three consecutive years of field experimentation in Mali, Striga reproduction was studied for 10 sorghum genotypes at infestation levels ranging from 30 000 to 200 000 seeds m^?2. Host resistance was identified as an important determinant of Striga reproduction, with the most resistant genotypes (N13, IS9830 and SRN39) reducing Striga reproduction by 70–93% compared with the most susceptible genotype (CK60‐B). Seedbank density had a significant effect on Striga seed production. Higher seedbank density resulted in more Striga plants, which led to increased intra‐specific competition and consequently a reduced level of reproduction per plant. For the most susceptible sorghum genotypes, density dependence also occurred in the earlier belowground stages. Striga reproduction continued beyond harvest. At the high infestation level just 8% of the total reproduction was realised after harvest, whereas at the low infestation level 39% was attained after harvest. Even though host‐plant genotype plays a significant role in Striga reproduction, calculations indicated that only at very low infestation levels the use of the most resistant genotype was able to lower the Striga seedbank.