向日葵列当生物学特性及抗列当向日葵分子育种研究进展

向日葵是世界各国重要的油料作物,也是供食用的重要经济作物。向日葵列当是危害向日葵生长极为严重的一类草害,在全球均有发生,且列当种子生活力极强,难以根除。向日葵列当研究和抗列当向日葵育种是农业科研人员研究的重点。本文综述了向日葵列当的生物学特性、寄生与传播特点、为害症状、预防和控制措施,以及近来年抗列当向日葵分子育种研究进展,并对抗列当向日葵育种研究发展前景作分析和展望。     

列当王国探奇

被子植物中的列当科,全是寄生草本植物,有15属150多种,中国有9属40多种。该科植物形态奇特,生活习性引人入胜,一些药用种类久已闻名。 列当小王国 列当(Orobanche coerulescens)是列当科的习见种之一,在我国北方分布较广,但以往见到的多是零星生长的少量植株。一年夏天,在山西五台山下的一片撂荒地田,笔者意外地见     

不同列当抗性的向日葵品种根系分泌物对向日葵列当种子萌发的影响

以对向日葵列当抗性不同的向日葵幼苗为材料,利用水培法收集其根系分泌物,经二氯甲烷萃取后结合硅胶柱色谱分离法,研究不同列当抗性的向日葵品种根系分泌物对向日葵列当种子萌发的影响。结果表明:(1)在一定浓度范围内,向日葵根系分泌物对向日葵列当种子萌发具有低浓度促进、高浓度抑制的作用;二氯甲烷萃取的向日葵根系分泌物浓度为0.1mg/mL时,其向日葵列当萌发率高于其他浓度的作用。(2)‘星火大白边’和‘白葵杂9号’根系分泌物提取物作用下向日葵列当萌发率最高分别为46.92%和43.88%,显著高于免疫品种‘MGS’根系分泌物提取物作用下列当的萌发率(27.81%)。(3)根系分泌物提取物正己烷-乙酸乙酯1∶1洗脱组分的萌发刺激活性显著高于其他组分,‘星火大白边’、‘白葵杂9号’和‘MGS’的该组分作用下向日葵列当最高萌发率分别为49.90%、45.66%和30.00%。(4)‘MGS’的根系分泌物提取物与GR24或脱氢木香内酯共同作用时种子萌发率显著低于GR24或脱氢木香内酯的单独作用,说明‘MGS’根系分泌物中可能含有抑制GR24或脱氢木香内酯刺激向日葵列当种子萌发的物质,使其对向日葵列当表现出免疫特性。     

土壤条件对向日葵列当寄生的影响

以高感列当的向日葵品种‘龙食葵2号’为试验材料,采用室内接种列当温室培养的方法,研究土壤条件对向日葵列当寄生与生长的影响。结果表明：（1）土壤质地对向日葵列当的寄生具有显著影响,培养50 d时,沙壤土、壤土、粘土中列当的出土数分别为 32.2、19.5和4.6个,平均单株寄生列当数量分别为38.8、24.0和8.2个,说明通透性好的沙壤土有利于列当瘤的形成与生长。（2）在沙壤土条件下,随着处理温度的上升,列当的出土数量和寄生数量均呈先升高后降低的趋势;当培养温度为25 ℃和30 ℃时,向日葵列当有较高的寄生数量与生长速度,列当出土数和平均单株寄生列当数量分别为30.4、31.6和39.8、39.4个。（3）在沙壤土条件下,随着土壤湿度的增加,列当的出土数量和寄生数量呈先升高后降低的变化趋势,并在土壤湿度为60%和70%达到较高值,列当出土数和平均单株寄生列当数量分别为32.6、30.9和39.9、40.9个,而在高湿条件下列当的寄生数量显著减少。（4）在沙壤土条件下,随着土壤pH值的增加,列当的出土数量和寄生数量也呈先升高后降低的变化趋势,并均在土壤pH值为8时达到最大值,列当出土数和平均单株寄生列当数量分别为47.5和51.1个,即列当在偏碱性的土壤环境中有着较高的寄生数量。研究发现,当土壤类型为沙壤土、土壤温度为25~30 ℃、土壤湿度为60%~70%、土壤 pH为8时有利于向日葵列当的寄生与生长;在实际生产中应选择中性或偏酸性的非沙壤土地种植向日葵,并注意调整播期和控制田间灌水量,以降低向日葵列当的危害。     

向日葵对列当寄生的生理响应及关键代谢通路分析

【目的】分析向日葵对列当寄生的生理响应及其相关代谢通路,揭示向日葵对列当寄生的抗性分子机制。【方法】以抗列当向日葵材料(s41)和感列当向日葵材料(s26)为供试材料进行接种列当处理,对正常生长(s41_CK,s26_CK)和列当寄生胁迫(s41,s26)条件下的4个样本进行转录组测序分析,同时对部分关键代谢物质进行生理验证分析。【结果】(1)在列当接种条件下,向日葵高感列当材料的植株高度和根量明显受到抑制,而且已有部分列当出土,而高抗列当材料无明显变化,仅有极少数列当寄生。(2)通过对样品间进行差异基因筛选,得到差异表达基因(DEGs)数量分别为6 362,6 609,5 490,5 469个。GO分析表明,s26-s41、s26_CK-s41_CK、s26-s26_CK、s41-s41_CK 4个比较组参与生物过程的差异基因数量较多,参与细胞组分的差异基因数量次之,参与分子功能的较少;KEGG分析表明列当寄生对向日葵的植物信号转导、次生代谢物的生物合成、脂质代谢、...     

中国食用向日葵育种国产化历程及研究进展

中国食用向日葵育种近40年来取得了巨大成就,实现了从自留农家种、选育常规品种、引进国外杂交种以及培育国产杂交种4 次品种更新。中国向日葵种植面积基本保持在59万hm²左右,种植区主要集中在北方10个省区,其中内蒙古自治区种植面积约占全国总面积的四分之三,其次是新疆。中国以食用向日葵种植为主,约占总面积的95%以上,是世界上食用向日葵主要生产国。食用向日葵主要用于休闲嗑食,除高产外,商品外观品质(粒型、色泽、口感等)仍将是当今主要育种目标。另外,向日葵生产区因长期连作种植,病原菌、列当等生物逆境因素与向日葵发生协同进化,加之食用向日葵遗传背景狭窄,病虫草害日趋严重。该文通过对中国食用向日葵育种历程进行回顾,尤其对国内外有关向日葵抗病育种研究进展进行综述,为解析食用向日葵抗性育种机制以及前瞻性抗性品种储备提供重要的理论与技术支持,以推动食用向日葵产业的高效优质发展。     

列当寄生对不同品种向日葵幼苗生长及抗氧化酶活性的影响

采用连作重茬含向日葵列当的土壤,进行不同列当抗性向日葵品种苗期盆栽试验,调查播种后第30、50和70天向日葵幼苗根系列当的寄生率、幼苗地上部和地下部干物质量及株高,并测定其叶片丙二醛(MDA)、苯丙氨酸解氨酶(PAL)、超氧化物歧化酶(SOD)、过氧化物酶(POD)和过氧化氢酶(CAT)活性。结果表明:不同向日葵品种苗期对列当抗性明显不同,其中油葵品种‘T012244’和‘MGS’对向日葵列当寄生免疫,‘S31’和‘白杂9号’为高抗,而‘星火大白边’和‘白杂6号’为敏感。列当寄生后明显抑制敏感品种向日葵幼苗生长,而对免疫品种和高抗品种的影响较小。列当寄生后的敏感性品种‘星火大白边’幼苗叶片MDA含量迅速上升,PAL活性也大幅度增加,列当对幼苗生长伤害加重;而苗期免疫品种"T012244"和高抗品种"S31"MDA含量下降,PAL活性则变化不大。随接种时间延长,不同抗性向日葵品种苗期SOD、POD和CAT活性均呈现先升高后降低的变化趋势,且免疫品种的保护酶活性变化幅度大于敏感品种,说明列当寄生对敏感向日葵品种造成了逆境胁迫,诱导其幼苗叶片保护酶系统产生防御响应。     

Streptomyces rochei D74菌剂对向日葵、列当及其根际微生物的影响北大核心

【目的】研究娄彻氏链霉菌(Streptomyces rochei)D74菌剂促进向日葵生长和抑制列当寄生的作用,及其对根际微生物的影响。【方法】通过构建向日葵-列当-S.rochei D74共培养体系,观察D74菌株对列当寄生的影响;采用田间小区试验,测定D74菌剂接种对向日葵植物生理指标和列当出土数的影响;测定向日葵产量指标,研究D74菌剂对向日葵籽粒品质及产量的影响;利用16S rRNA基因高通量测序技术和传统微生物培养方法,分析添加D74菌剂对向日葵根际微生物群落结构的影响。【结果】D74菌剂能够促进向日葵生长[茎秆重和花盘重分别显著增加(P<0.05)37%-40%和21%-37%],同时抑制28%-46%(P<0.05)的列当出土;还可以提高向日葵籽粒中粗蛋白含量5%-9%、大粒占比约66%(P<0.05)和百粒重8%-18%,并且通过产量测定表明,D74菌剂可增产约30%(P<0.05),这对提高农民经济收益具有深远意义。D74菌剂对向日葵根际微生物群落具有显著影响,调节了群落有益微生物数量。【结论】D74菌剂可以显著降低向日葵根部列当寄生病害的发生,促进花盘生长,使籽粒饱满,具有实际应用价值和推广意义。     

盐生肉苁蓉寄主植物种类鉴定及分布区域分析

盐生肉苁蓉（Cistanche salsa (C. A. Mey.) G. Beck）为列当科肉苁蓉属寄生性草本植物,为目前已知寄主植物种类最多的肉苁蓉属物种,也是我国西北地区传统常用中药材。为探明我国盐生肉苁蓉寄主植物种类及分布区域,本研究结合DNA条形码鉴定和形态鉴别技术,对采集到的66份带有寄主根的盐生肉苁蓉样品进行分布区域分析和寄主植物种类鉴定。结果表明盐生肉苁蓉在新疆及宁夏等地区海拔400～2000 m的区域分布广泛。新疆塔城地区裕民县盐生肉苁蓉寄主植物为菊科绢蒿属聚头绢蒿（Seriphidium compactum (Fisch. ex DC.) Poljakov）和苋科滨藜属疣苞滨藜（Atriplex verrucifera Marsch. von Bieb.）,塔城市盐生肉苁蓉寄主植物为苋科盐爪爪属里海盐爪爪（Kalidium cuspidatum Ung.-Sternb.）、疣苞滨藜和驼绒藜属驼绒藜（Krascheninnikovia ceratoides (L.) Gueldenst.）,该地区优势寄主植物为疣苞滨藜。新疆阿勒泰地区哈巴河县寄主植物为聚头绢蒿、苋科驼...     

新疆荒漠绿洲生态区苹果蠹蛾生物学、生态学和防治技术研究与应用进展

苹果蠹蛾是我国重要的检疫害虫,自1953首次在新疆库尔勒地区发现以来,疫情保持不断扩张的趋势,目前已经在我国新疆、甘肃、宁夏、内蒙古、黑龙江、辽宁和吉林7个省区发现,对我国苹果产业构成了严重威胁。新疆在苹果蠹蛾生物学、生态学等研究领域起步较早,开展了大量的相关研究和应用工作。本文通过对新疆苹果蠹蛾相关文献的搜集、整理和分析,回顾和总结了新疆绿洲荒漠生态区苹果蠹蛾生物学、生态学和综合防治技术等领域的研究和应用进展,以期对今后该虫的研究和防治提供参考。     

Induction of seed germination in <Emphasis Type="Italic">Orobanche</Emphasis> spp. by extracts of traditional Chinese medicinal herbs

The co-evolution of Orobanche spp. and their hosts within the same environment has resulted in a high degree of adaptation and effective parasitism whereby the host releases parasite germination stimulants, which are likely to be unstable in the soil. Our objective was to investigate whether extracts from non-host plants, specifically, Chinese medicinal plants, could stimulate germination of Orobanche spp. Samples of 606 Chinese medicinal herb species were extracted with deionized water and methanol. The extracts were used to induce germination of three Orobanche species; Orobanche minor, Orobanche cumana, and Orobanche aegyptiaca. O. minor exhibited a wide range of germination responses to the various herbal extracts. O. cumana and O. aegyptiaca exhibited an intermediate germination response to the herbal extracts. O. minor, which has a narrow host spectrum, showed higher germination rates in response to different herbal extracts compared with those of O. cumana and O. aegyptiaca, which have a broader host spectrum. Methanolic extracts of many Chinese herbal species effectively stimulated seed germination among the Orobanche spp., even though they were not the typical hosts. The effective herbs represent interesting examples of potential trap crops. Different countries can also screen extracts from indigenous herbaceous plants for their ability to induce germination of Orobanche spp. seeds. The use of such species as trap plants could diminish the global soil seed bank of Orobanche.     

Isolation, Identification, and Activity of Mycoherbicidal Pathogens from Juvenile Broomrape Plants

Although there are reports of isolation of mycoherbicidal pathogens attacking the widespread broomrapes (Orobanche spp.) that parasitize legumes and vegetables, none is in use or available. This is despite there being no good method of controlling broomrapes in most crops other than by preplant fumigation with methyl bromide. Two highly parasitic fungi, Fusarium arthrosporioides strain E4a (CNCM I-164) and F. oxysporum strain E1d (CNCM I-1622), were isolated from nearly 100 organisms found on diseased, juvenile, emerging Orobanche flower stalks. A near-axenic polyethylene envelope system for culturing broomrape on tomato roots was used to ascertain pathogenicity of these strains. Both organisms fulfilled Koch's postulates for being primary pathogens. Their DNAs were analyzed and fingerprinted by restriction fragment length polymorphism and random amplified polymorphic DNA, showing that they are indeed different from each other and from many other Fusarium spp. and other formae speciales of F. oxysporum including a strain that attacks O. cumana on sunflowers. Both strains infect O. aegyptiaca, O. cernua, and O. ramosa, but not O. cumana. They did not infect any of the vegetable and legume crops tested and thus seem specific to Orobanche. Tomato plant roots dipped into a fungal spore and mycelial suspension and planted in broomrape-infested soil were protected for 6 weeks, as were tomato transplants in pot experiments. About 90% control was also achieved by posttransplant soil drench with fungal suspensions in pot experiments. These pathogens may be effective as seed, transplant, or soil-drench treatments of high-value vegetable and other crops.     

Recent advances in the biocontrol of Orobanche (broomrape) species

Parasitic broomrapes (Orobanche spp.) are majoruncontrolled weeds in the Mediterranean regions of Europe and the NearEast causing major losses to vegetable, grain legume, and sunflowercrops. Selective herbicides alone cannot provide persistent, season-longcontrol of these parasites, and much methyl bromide is used for theircontrol, where affordable. Thus they are excellent targets forbiocontrol. The recent progress by the COST 816 Orobancheworking group in this area is reviewed herein. Natural infestation bythe fly Phytomyza orobanchia of seed capsules of Orobanchecrenata parasitising faba bean halved Orobanche seedproduction while inundative releases of adults reduced it to 5%of viable seeds. The fungi Fusarium arthrosporioides E4a andF. oxysporum E1d, as well as strains of bacteria were isolatedfrom diseased, juvenile, Orobanche flower stalks. They arepathogenic to O. aegyptiaca, O. crenata and O. ramosaon most vegetable crops. A F. oxysporum f. sp.orthoceras was specifically pathogenic to O. cumana onsunflowers. All were used in various experiments with a modicum ofsuccess. Methods were developed to formulate isolated mycelia, whichcould eventually allow the use of transgenic hypervirulent pathogens inasporogenic (deletion) mutants (as a failsafe against spread).Mycotoxins were also isolated from different Fusarium and otherfungal species that kill Orobanche, and are being consideredfor direct use, or to augment other strategies. All threeFusarium spp. used have been transformed with gusand/or gfp genes allowing tracing their movement in theenvironment, and opening the way to future transformations tohypervirulence.     

Use of random amplified polymorphic DNA (RAPD) markers in the study of the parasitic weed Orobanche

Despite the tremendous economic impact of broomrapes (Orobanche spp.) on agriculture in many countries little is known of the pattern of genetic variation within this group of parasitic weeds. The present paper describes the use of RAPD markers for the study of five Orobanche species in agricultural fields in Israel. Pronounced genetic differentiation was found between the species, and RAPD markers were raised for the identification of each of them. Southern-hybridization patterns of RAPD products of the various species were used to confirm the interpretation. The same markers were valid both for broomrapes collected in agricultural fields and for those collected in natural habitats. The validity of the markers found for O. cumana and O. crenata was confirmed on plants of the same species that were collected in Spain. Parsimony analysis of 86 RAPD characters produced a tree that clearly distinguishes between the five studied Orobanche species, separates the two Orobanche species belonging to sect. Trionychon from those belonging to sect. Osproleon, and supports the separation of O. cumana from O. cernua and of O. aegyptiaca from O. ramosa.     

Biological control of Orobanche spp. with Phytomyza orobanchia, a review

This review summarises theavailable information on the biocontrol agentPhytomyza orobanchia Kalt. (Diptera, Agromyzidae). It gives an overview of a rareexample of weed biocontrol using insects in aninundative approach. A high diversity ofphytophagous insects has been collected onparasitic weeds of the genus Orobanche(Orobanchaceae). For biological control of Orobanche spp., only insects like P. orobanchia whose host range is restricted toOrobanche spp. are of interest. Of the140 Orobanche spp. described in total,P. orobanchia is reported from 21 species. The larvae of P. orobanchia minein Orobanche shoots and capsules. As aconsequence, a natural reduction of Orobancheseed production by 30 to almost 80%has been reported from different countries. Theefficacy of P. orobanchia under naturalconditions is limited by low temperatures, cultural practices and natural enemies. Tostrengthen the natural population and itsimpact, inundative releases of P. orobanchia adults at the beginning of Orobanche emergence have to be undertaken. Different methods for the application of P. orobanchia in biocontrol of Orobanche spp. have been developed in theformer Soviet Union. Releases of 500 to 1000adults/ha resulted in a reduction of up to 96%of the Orobanche seed production.However, due to the 10 to 15 year longevity ofOrobanche seeds, further infestationswill occur in the following cropping seasons.Consequently, releases of P. orobanchiahave to be repeated continuously over severalyears to reduce the infestation to a tolerablelevel.     

Die Chromosomenzahlen der in Mitteleuropa vorkommenden Arten vonOrobanche sect.Orobanche

The chromosome numbers of five species ofOrobanche sect.Orobanche (O. alsatica, O. laserpitii-sileris, O. loricata, O. salviae, O. teucrii) are reported for the first time and previous counts could be verified in ten other species. Now the chromosome numbers of all species of sect.Orobanche occurring in Central Europe are known: they are diploid (2n = 38) with the exception ofO. gracilis (tetra- and hexaploid, aneusomatic).

     

Colonization and predation of thrips (Thysanoptera: Thripidae) by Orius spp. (Heteroptera: Anthocoridae) in sweet pepper greenhouses in Northwest Italy

Frankliniella occidentalis (Pergande) and Thrips tabaci Lindeman (Thysanoptera: Thripidae) are major pests of sweet pepper for direct damage and tospovirus transmission. To control their infestations, Orius laevigatus (Fieber) (Heteroptera: Anthocoridae) is produced by many commercial insectaries and widely used on IPM vegetable crops of Europe. This predator is naturally widespread along the Mediterranean and Atlantic coasts, and not in more continental areas, where other Orius spp. are more common. Research was conducted in a continental area of Northwest Italy in 2002–2003 to assess the natural presence of anthocorids on pepper, and to compare their colonization and predatory ability with those of the species artificially introduced. Experiments were conducted in 12 sweet pepper greenhouses, in six of which O. laevigatus release was made. From late May to early October, thrips and anthocorids were sampled on pepper by collecting flowers; Orius spp. were also collected on neighboring wild flora. Independently of the releases, Orius specimens were found in all surveyed greenhouses, but O. niger Wolff, also captured on various wild plants, was the most abundant species. It naturally colonized crops from late June and proved to be the most efficient predator on sweet pepper in the surveyed area, if not disturbed by pesticide application. Contrarily, O. laevigatus was rarely found and only in the greenhouses in which it had been released. However its introduction resulted in thrips control before natural colonization by the native species occurred.     

二疣犀甲对油棕、椰子的风险性分析及防控策略

二疣犀甲是危害椰子、油棕等木本油料作物的重要害虫,本文从形态学、生物学、分布及危害等不同方面对该虫进行了阐述,并对其风险性进行了定性定量评价,结果表明:二疣犀甲的R值为2.0,属于高风险有害生物。同时从生物防治、物理防治、化学防治、农事操作及检验检疫技术的应用等方面制定了二疣犀甲的综合防控策略,为该虫的危害评价和防护技术的制定提供了参考。     

Host plant resistance against broomrapes (Orobanche spp.): defence reactions and mechanisms of resistance

Over 4000 species of angiosperms are able to directly invade and parasitise other plants, but only very few are weedy and parasitise cultivated plants. Together with the witchweeds (Striga spp.) and dodders (Cuscuta spp.), the broomrapes (Orobanche spp.) affect important crops causing complete yield loses with severe infestations. Genetic resistance to parasitisation remains as one of the most desirable components in an integrated control strategy. However, breeding for resistance is a difficult task and many aspects of the host/parasite interaction remain unknown. In the present work, we review the cytological and cytochemical studies investigating the mechanisms of resistance involved in the process leading to an incompatible host–parasite interaction. We have attempted to identify the main gaps and problems related to such studies with parasitic plants and present an in‐depth review of histochemical techniques used to assess mechanisms of resistance against these parasitic plants. Furthermore, we discuss future research directions and novel techniques and finally, how such techniques can be applied and incorporated within a breeding programme.     

The potential of Ulocladium botrytis for biological control of Orobanche spp.

A strain of Ulocladium botrytis isolated from diseased Orobanche crenata shoots caused disease on the parasitic weed in pathogenicity tests. The potential of the fungus to be developed as a mycoherbicide for Orobanche spp. was further investigated. Although the fungus significantly decreased O. crenata germination in vitro by 80%, it did not generally lead to a decreased number of O. crenata shoots or tubercles in inoculated root chambers or pots. However, the number of diseased or dead tubercles and underground shoots was significantly increased compared to the noninoculated treatments. Postemergence inoculation of O. crenata shoots with a conidial suspension resulted in the death of almost all inoculated plants 14 days after application under greenhouse conditions. In preliminary host-range studies, the pathogen caused disease on Orobanche cumana on sunflower whereas on Orobanche aegyptiaca shoots parasitizing tomato only minimal disease symptoms could be detected after postemergence inoculation. Based on the results of our investigations, we conclude that Ulocladium botrytis has only a limited potential to be used as a biocontrol agent against Orobanche spp.