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。     

New insights concerning the ecology and the biology of Rhamphicarpa fistulosa (Scrophulariaceae)

Rhamphicarpa fistulosa (Hochst.) Benth. is a parasitic member of the Scrophulariaceae occurring sporadically on rice and other cereal crops in tropical Africa. Relatively little is known about this parasite, and some aspects concerning its life cycle and its impact on crops are reported in the present paper. The size, the shape and the seedcoat ornamentation of R. fistulosa are described and compared with those of Striga spp. The seeds do not require a host root exudate to stimulate germination but do require light. As in the case of Striga spp., they show a period of dormancy of at least 6 months. When the parasite roots come into close contact with a host root, they swell and develop haustoria, which form a direct xylem connection between host and parasite. In vitro and pot culture experiments establish that R. fistulosa is a facultative parasite, as it is able to complete its life cycle without a host plant. However, plants growing under these conditions are much smaller and produce only a few seeds. Field observations indicate that in some places, R. fistulosa can destroy cereals. The agronomic hazard represented by this member of the Scrophulariaceae is discussed.     

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。     

Influence of Root Exudates on Orobanche Seed Germination*

Broomrape (Orobanche ramosa L.) is a common root parasite of solanaceous, leguminous and other crops grown in the semi-arid regions of the world. The seeds germinate when root exudates from host plants are released in their immediate vicinity (Lindley, 1853; Koch, 1887; Chabrolin, 1934). Brown et al. (1951a) reported that non-host plants, such as flax (Linum usitatissimum L.) may stimulate Orobanche seed germination without being parasitized. The stimulating properties of flax exudate were studied by Brown et al. (1951b). They reported that the stimulant was unstable in alkaline solutions, but moderately stable in weakly acidic media, which may indicate the presence of an acidic (lactone) grouping. Nash & Wilhelm (1960) reported that gibberellic acid in agar media stimulated O. ramosa seed germination. Abu- Shakra, Miah & Saghir (1970) found that pre-treatment of 0. ramosa seeds with 100 ppm of gibberellic acid followed by incubation on a flax-root diffusate agar medium gave a high (81·7%) germination. The purpose of this study was to collect root exudates from three species of plants cultured under three experimental systems, namely (a) germ-free, (b) glasshouse (non-sterile), and (c) growth chamber (hydroponic, initially aseptic), and to evaluate their biological activity as germination stimulants for O. ramosa seeds. The plants used were tomato (Lycopersicon esculentum Mill.), sorghum (Sorghum vulgare Pers.) and flax. Exudate from marigold (Tagetes erecta L.) also was collected from germ-free culture.     

Some compatible Rhizobium leguminosarum strains in peas decrease infections when parasitised by Orobanche crenata

Orobanche crenata (crenate broomrape) produces serious damage to many legume crops and particularly becomes a limiting factor for pea production in the Mediterranean basin. Nodulation effects on pea–broomrape relationships were studied using the commercial pea cultivar Douce de Provence and different Rhizobium strains using pot and Petri dish experiments. First, the benefit of bacterial inoculation on plant growth and efficiency of N incorporation were demonstrated for two isolates, P.SOM and P.1236. These isolates did not influence parasite germination induced by the artificial stimulant, GR24. In contrast, pea root inoculation with P.SOM and P.1236 isolates led to a reduced root infection by O. crenata , resulting from a lower Orobanche germination rate close to pea roots and a limited capacity of the parasitic seedlings to develop tubercles. Broomrape necrosis was observed both before and after parasite attachment to inoculated pea roots. Concomitantly, reduction in infection was accompanied by enhanced peroxidase activity and constantly high phenylalanine ammonia lyase activity in pea roots. These data suggest the involvement of these enzymes in pea resistance to crenate broomrape induced by the compatible rhizobia. Management of Orobanche via crop selection based on these enzyme systems is a viable option.     

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.     

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.     

Suicidal germination for parasitic weed control

Parasitic weeds of the genera Striga and Orobanche spp. cause severe yield losses in agriculture, especially in developing countries and the Mediterranean. Seeds of these weeds germinate by a chemical signal exuded by the roots of host plants. The radicle thus produced attaches to the root of the host plant, which can then supply nutrients to the parasite. There is an urgent need to control these weeds to ensure better agricultural production. The naturally occurring chemical signals are strigolactones (SLs), e.g. strigol and orobanchol. One option to control these weeds involves the use of SLs as suicidal germination agents, where germination takes place in the absence of a host. Owing to the lack of nutrients, the germinated seeds will die. The structure of natural SLs is too complex to allow multigram synthesis. Therefore, SL analogues are developed for this purpose. Examples are GR24 and Nijmegen‐1. In this paper, the SL analogues Nijmegen‐1 and Nijmegen‐1 Me were applied in the field as suicidal germination agents. Both SL analogues were formulated using an appropriate EC‐approved emulsifier (polyoxyethylene sorbitol hexaoleate) and applied to tobacco (Nicotiana tabacum L.) fields infested by Orobanche ramosa L. (hemp broomrape), following a strict protocol. Four out of 12 trials showed a reduction in broomrape of ≥95%, two trials were negative, two showed a moderate result, one was unclear and in three cases there was no Orobanche problem in the year of the trials. The trial plots were ca 2000 m²; half of that area was treated with stimulant emulsion, the other half was not treated. The optimal amount of stimulant was 6.25 g ha^?1. A preconditioning prior to the treatment was a prerequisite for a successful trial. In conclusion, the suicidal germination approach to reducing O. ramosa in tobacco fields using formulated SL analogues was successful. Two other options for weed control are discussed: deactivation of stimulants prior to action and biocontrol by Fusarium oxysporum. © 2016 Society of Chemical Industry     

Possible involvement of gibberellins and ethylene in Orobanche ramosa germination

Summary Six inhibitors acting at different stages of gibberellin biosynthesis, applied during conditioning of Orobanche ramosa seeds, reduced subsequent germination in the presence of GR₂₄ (a strigol analogue). Ethylene seems to be involved in the induction of germination of conditioned seeds by GR₂₄, as inhibitors of its synthesis or action, applied to conditioned seeds, also strongly reduced induction of germination by GR₂₄. Exogenous ethylene did not induce germination of conditioned seeds, but 2-chloroethylphosphonic acid was able to do so. When inhibitors of gibberellin biosynthesis were applied to conditioned seeds in the presence of GR₂₄, they inhibited germination. These same inhibitors also strongly inhibited germination of conditioned Striga hermonthica seeds in response to GR₂₄; this inhibitory effect was reversed by the addition of 2-chloroethylphosphonic acid. The effect of these inhibitors on S. hermonthica , in which ethylene is a necessary mediator of germination induction by GR₂₄, strongly suggests that ethylene synthesis is also required for the induction of O. ramosa seed germination by GR₂₄. These growth regulators, which inhibit the two steps of germination in O. ramosa , could be useful for the development of methods for early season control of this parasite.     

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.     

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。     

Possible peroxidase functions in the interaction between the parasitic plant, Orobanche aegyptiaca, and its host, Arabidopsis thaliana

The mechanism by which the flowering holoparasitic plant, Orobanche aegyptiaca , infects its host without evoking a defence mechanism is still poorly understood. In this work, we studied several mechanisms used by phytopathogenic fungi. We focussed on the possible role of peroxidases during O. aegyptiaca penetration into the roots of Arabidopsis thaliana . A convenient experimental system for studying the interaction under sterile conditions was developed. The formation of extracellular reactive oxygen species (ROS) was detected at the interaction site before, during, and after the parasite penetrated into the host. These extracellular ROS probably originated from the parasite. However, no intracellular ROS could be detected at the site of the interaction. Peroxidase activity was observed mainly at the apex of the root of the parasite and in the adventitious roots of the tubercle. Benzhydroxamic acid, a peroxidase inhibitor, was used to probe the possible role of peroxidase in the infection process. Peroxidase activity was observed in the root apex and adventitious roots of O. aegyptiaca, but no evidence was found for its participation in the actual infection process. Peroxidase activity was also found in the later stages of the interaction between the host and the parasite. We propose that peroxidases could have a role in generating extracellular ROS for loosening the cell wall of the host in order to facilitate penetration. Alternatively, the ROS could act in facilitating the root elongation of the parasite.     

Germination response of Orobanche seeds subjected to conditioning temperature, water potential and growth regulator treatments

Broomrapes (Orobanche spp.) are parasitic weeds that cause significant losses of crop yield. Experiments were conducted to investigate the seed response to the artificial germination stimulant GR₂₄ in three species of Orobanche subjected to preconditioning under various temperatures, water potentials and with plant growth regulators. The highest germination percentages were observed in Orobanche ramosa, Orobanche aegyptiaca and Orobanche minor seeds conditioned at 18°C for 7 days followed by germination stimulation at 18°C. With the increase of the conditioning period (7, 14, 21 and 28 days), the germination percentage of O. ramosa and O. aegyptiaca progressively decreased. When conditioned at −2 MPa, the germination percentage was lower than at 0 and −1 MPa, especially at 13 and 28°C. Orobanche minor seeds could retain relatively high germination if conditioned at 18, 23 or 28°C, even after significantly extended conditioning periods (up to 84 days). GA₃ (30–100 mg L⁻¹), norflurazon and fluridone (10–100 mg L⁻¹), and brassinolide (0.5–1.0 mg L⁻¹) increased seed germination, while 0.01 mg L⁻¹ uniconazole significantly reduced germination rates of all three Orobanche spp. The promotional effects of GA₃ and norflurazon and the inhibitory effect of uniconazole were evident, even when they were treated for 3 days. Germination of Orobanche seeds was much lower when the unconditioned seeds were directly exposed to GR₂₄ at 10⁻⁶ m . This early GR₂₄-induced inhibition was however alleviated or even eliminated by the inclusion of GA₃ or norflurazon (10–50 mg L⁻¹) in the conditioning medium. On the contrary, the inclusion of uniconazole increased the inhibitory effect of GR₂₄, particularly in the case of O. ramosa.     

Germination of <Emphasis Type="Italic">Plasmodiophora brassicae</Emphasis> resting spores stimulated by a non-host plant

Plant-induced germination of Plasmodiophora brassicae resting spores was studied in a laboratory experiment. Spore reaction was analysed in nutrient solution with exudates from growing roots of different plant species – one host plant (Brassica rapa var. pekinensis) and four non-host plants (Lolium perenne, Allium porrum, Secale cereale and Trifolium pratense) – and in controls with distilled water and nutrient solution. It was found that root exudates from L. perenne stimulated spore germination more than exudates from the other plants, including those from the host plant. The effect could not be explained by differences in the nutritional composition of the solutions due to differential uptake of the plant species, or by differences in root activity, measured as exudation of soluble sugars. This is the first time such a separation of factors has been done in analysing the influence of plants on P. brassicae germination. Although stimulation of P. brassicae resting spore germination is not restricted to the presence of host plants, it seems to vary depending on the plant species.     

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.     

Methyl jasmonate and methyl salicylate, but not cis-jasmone, evoke defenses against infection of Arabidopsis thaliana by Orobanche aegyptiaca

The mechanism by which some plant species develop resistance to the root parasite, broomrape ( Orobanche aegyptiaca ), is still not clear. Resistance to other pathogens can be induced by methyl jasmonate and systemic acquired resistance can be induced by treatment with salicylic acid, while cis -jasmone can act as a signaling molecule in plant–insect interactions. The three compounds studied, methyl jasmonate, cis -jasmone, and methyl salicylate, were applied to Arabidopsis thaliana seedlings that were then transferred to Nunc cell culture plates and exposed to the germinating seeds of O. aegyptiaca . The number of infections of the roots of single seedlings of A. thaliana was then quantified. Exposure for 24 h to very low concentrations of methyl jasmonate or methyl salicylate, which were then removed, effectively induced resistance to infection of A. thaliana by O. aegyptiaca , reducing attachment and tubercle formation by 90%. cis -Jasmone was far less effective in inducing a similar resistance to infection. These results support the view that methyl jasmonate can induce almost full resistance to infection by broomrape. The fact that such resistance is not observed under normal conditions of infection supports the idea that the root parasite does not evoke the full defensive response in the host plant.     

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.     

Presowing Hardening of the Host with Phenolic Acids Reduces Induction of Seed Germination in the Root Parasite Striga asiatica

Abstract

Striga asiatica (L.) Kuntze, a root parasite, causes severe loss of yield in sorghum and several other crops. The seeds of the parasite are induced to germinate by a stimulant in the host root exudate. Presowing hardening of the host with vanillic acid, caffeic acid and ferulic acid (25 ppm) reduces the induction of seed germination in the parasite by the host root exudate. The treatment causes a slight improvement in the dry matter production in the host and in addition, increases the phenolics level in the host root exudate. The latter effect might be responsible for reducing germination in Striga. If the treatment remains effective under field conditions also, it reduces significantly the incidence of Striga in cultivated fields.     

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 .