Intraspecific seasonal variation of dormancy and mortality of Phelipanche ramosa seeds

Phelipanche ramosa (Branched broomrape) is an obligate root parasitic plant that is a major pest of oilseed rape in France. Knowledge on seed viability and dormancy under field conditions is crucial to understand how to control P. ramosa, but is as yet unknown. Our study aimed to quantify these processes with a 2‐year seed burial experiment. Two genetically distinct populations of P. ramosa were studied, collected on winter oilseed rape (population O) and hemp (population H). Seed mortality was very low in both populations (4–7% per year). Although obligate parasitic seeds are assumed to germinate only after exposure to germination stimulants from host root exudates, a high proportion of population H seeds germinated spontaneously (up to 90%). Seeds of both populations displayed seasonal dormancy, with timing and magnitude depending on the population. Dormancy was low at the time each native host crop is usually sown. Populations differed in germination dynamics, with seeds of population H germinating faster. The difference in behaviour that we observed between populations is consistent with reported adaptations of pathovars to their preferred hosts. The results indicate that the parasitic plant management requires targeting at the populations concerned. For example, delayed sowing is more promising against population O than against population H.     

Genetic differentiation and host preference reveal non‐exclusive host races in the generalist parasitic weed Phelipanche ramosa

We developed 20 microsatellite markers to genotype over 100 populations of the parasitic weed Phelipanche ramosa, which covers a wide host crop and geographic range. A representative core collection of 15 populations was also used in cross‐infestation assays to study host preference during germination, attachment and shoot formation. We observed low genetic differentiation within most of the populations, but high genetic differentiation between populations partitioned into 3 genetic groups with different host preferences and geographic distributions. Genetic group 1 is detected exclusively in western France and on various host crops, notably winter oilseed rape (WOSR) and not hemp. Cross‐infection assays confirmed its incompatibility with hemp and showed its preference for WOSR and tobacco in terms of germination and attachment success. The group 2 populations share a large geographic distribution in France and Europe, low germination success with WOSR and high germination success, attachment success and shoot formation with hemp, tobacco or tomato. The subclades 2a and 2b include most of the French populations in hemp crops in eastern France and in tobacco fields in several European countries respectively. The genetic analyses revealed the potential of the three groups to increase their geographic range in the future. Intermediate genetic groups showed higher intrapopulation diversity and represent potential stocks for new host race emergence. Those findings argue in favour of the existence of host races in P. ramosa and should be considered for appropriate management strategies, notably in breeding programmes for resistance against this parasitic weed.     

根寄生杂草种子萌发剂概述

独脚金属（Striga spp.）及列当属（Orobanche spp.和Phelipanchce spp.）杂草是世界范围内对农业生产危害严重的根寄生杂草,主要依靠寄生茄科、豆科、谷类等作物获得水分和营养。根寄生杂草种子只有在合适的萌发诱导物（如独脚金内酯）刺激下才能萌发。文章从根寄生杂草的分类、分布及危害,种子萌发剂类型及适用范围,种子萌发剂独脚金内酯的生物合成及分离鉴定,根寄生杂草种子萌发诱导剂的作用机制以及种子萌发过程中激素间协同调控作用等方面进行了综述,并展望了种子萌发剂设计筛选及其在根寄生杂草化学防控中的潜在应用前景。     

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.     

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.     

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.     

Emerging technologies for the chemical control of root parasitic weeds

Parasitic plants in the Orobanchaceae family include devastating weed species, such as Striga, Orobanche, and Phelipanche, which parasitize major crops, drastically reduces crop yields and cause economic losses of over a billion US dollars worldwide. Advances in basic research on molecular and cellular processes responsible for parasitic relationships has now achieved steady progress through advances in genome analysis, biochemical analysis and structural biology. On the basis of these advances it is now possible to develop chemicals that control parasitism and reduce agricultural damage. In this review we summarized the recent development of chemicals that can control each step of parasitism from strigolactone biosynthesis in host plants to haustorium formation.     

Effects of individual plant phenology on dormancy of Rumex obtusifolius seeds at dispersal

In 1998–2000, the relationship was investigated between the time of bolting of shoots and the percentage of germinability in seeds from individual plants of Rumex obtusifolius . Each year, the date of bolting of shoots was recorded in 30–108 plants, which were then grown intact until maturation, and their seeds were collected before dispersal. The proportion of germinable seeds increased with the date on which the shoots bolted. The timing of shoot bolting and seed germinability for individual plants persisted over the years of observation. To discriminate between the effect of mother plant and timing of seed maturation, shoot bolting of another 40 (1998) and 50 (1999) plants was recorded, and their shoots were then cut in late June The shoots of cut plants regrew and matured nearly simultaneously, later than in intact plants, whereas overall seed germinability decreased. In individual plants, the seed germinability was still significantly correlated with the date when plants originally bolted.     

Early stages of infection of maize (Zea mays) and Pennisetum setosum roots by the parasitic plant Striga hermonthica

Infection of young roots of maize (Zea mays L.) by the parasitic plant Striga hermonthica (Del.) Benth. was examined. Attachment to and penetration of roots occurred within 1–2 days after inoculation. Subsequent growth through the cortex to the host stele and proliferation of parasite xylem tissue was commonly completed by 3–4 days after inoculation. Histochemical staining showed that young maize roots do not contain major wall-thickening components. However, an increase in cell wall fluorescence and endodermal cell wall thickness was often seen at the site of infection and in the surrounding maize root tissue at 3 days after inoculation. This host response was variable and did not prevent rapid and successful penetration by the parasite. In contrast, uninfected roots of Pennisetum setosum (Sw.) L. Rich., a species resistant to S. hermonthica, had substantial thickening of the inner endodermal cell walls and exhibited further cell wall thickening at the stele upon infection. Examination of infections on both hosts demonstrated the presence of autofluorescent material at the host-parasite interface. This material was thicker and more extensive at the P. setosum-S. hermonthica interface than at the maize-S. hermonthica interface, and contained polyphenols and lignin. Examination of the host-parasite xylem connections in maize revealed substantial invasion of the host stele by both parenchyma and tracheary elements. In a few cases of P. setosum infection, parasite cells entered the stele; however, this did not lead to successful establishment of the parasite.     

7种激素对蒺藜状苜蓿种子发芽的影响

本实验旨在研究蒺藜状苜蓿种子发芽对外源激素的响应。使用6-BA、GA3、ABA、NAA、乙烯利I、AA和IBA共7种激素,每一激素各设0.1、1.0、1.5、2.0、5、10、201、000μmol/L共8种浓度梯度,浸湿滤纸发芽床,置50粒蒺藜状苜蓿种子进行发芽试验,以蒸馏水处理为对照,设3个重复,统计发芽率、发芽势、发芽指数和发芽天数。结果显示:外源激素20μmol/L 6-BA、GA3、乙烯利和5μmol/L IAAI、BA促进蒺藜状苜蓿种子发芽,NAA和ABA抑制蒺藜状苜蓿种子发芽,ABA抑制能力较强。     

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.     

Impacts of snow on seed germination are independent of seed traits and plant ecological characteristics in a temperate desert of Central Asia

Seed germination profoundly impacts plant community composition within the plant life cycle. Snow is an important source of water for seed germination in the temperate deserts of Central Asia. Understanding how seed germination responds to variations in snow cover in relation to seed traits and plant ecological characteristics can help predict plant community sustainability and stability in Central Asia under a scenario climate change. This study investigated the seed germination of 35 plant species common to the Gurbantunggut Desert in Central Asia under the three snow treatments: (1) snow addition; (2) ambient snow; and (3) snow removal. Two-way analysis of variance (ANOVA) tests were performed to assess interactions among the impacts of snow treatments, seed traits and plant ecological characteristics on seed germination. Phylogenetic generalized least-squares (PGLS) model was used to test the relationships between seed traits and seed germination. The results demonstrated that snow variations had no significant impacts on seed germination overall. Seed germination under the snow addition treatment was similar with that under the ambient snow treatment, irrespective of seed traits and plant ecological characteristics. Snow removal only had negative impacts on seed germination for certain groups of seed traits and plant ecological characteristics. Seed mass positively affected seed germination, showing a linear increase of arcsin square root-transformed seed germination with log-transformed seed mass. Seed shape also profoundly impacted seed germination, with a higher germination percentage for elongated and flat seeds. Seed germination differed under different plant life forms, with semi-shrub species showing a significantly higher germination percentage. Most importantly, although snow treatments, seed traits and plant ecological characteristics had no interactive effects on seed germination overall, some negative impacts from the snow removal treatment were detected when seeds were categorized on the basis of seed mass and shape. This result suggests that variations of snow cover may change plant community composition in this temperate desert due to their impacts on seed germination.     

Changes in germinability, dormancy and viability of Amaranthus retroflexus as affected by depth and duration of burial

Changes in dormancy and viability of Amaranthus retroflexus seeds were examined after placement in pots that were buried in the field. Seeds were placed in woven nylon envelopes on the soil surface or buried at depths of 2.5, 5 or 10 cm. After 1, 3, 6, 9 and 12 months seeds were exhumed and their germinability was tested to assess changes in dormancy and viability. Depletion of seed stocks placed on the soil surface was partly because of in situ germination that did not exceed 21% and did not vary significantly over the 12-month study period. Less germination of buried seeds occurred in situ , and seeds that did not germinate appeared to acquire dormancy. Exhumed seeds germinated readily; germinability was linearly related to the depth of burial, with those retrieved from the surface germinating least. Cyclical changes in germinability occurred during the 12-month burial period, but this influence was identical for seeds buried at all depths. Germinability was greatest after periods with the lowest mean monthly temperatures and least during the hottest periods. The stimulation of remaining ungerminated seeds exhumed at each period, by the addition of ethephon to the germination medium, provided further evidence of a seasonal acquisition of dormancy, and it was concluded that other unknown factors besides cyclical changes in seasonal temperature were responsible. Irrespective of placement, all seeds lost viability at an exponential rate over time. However, the decline was most rapid for those placed on the surface, whereas the loss in viability became less with increased depth of burial. Possible explanations for this adaptation of enhanced survival when buried are discussed.     

Corynelia uberata as a threat to regeneration of Podocarpus falcatus in Ethiopian forests: spatial pattern and temporal progress of the disease and germination studies

The order Coryneliales includes several fungi such as Corynelia spp. that are pathogenic to trees in the Podocarpaceae. The aim of this study was to assess the spatial pattern and temporal progress of disease caused by Corynelia uberata on Podocarpus falcatus in Ethiopian forests and to evaluate the germination potential of seed retrieved from fruit infected by C. uberata. Corynelia uberata was found on leaves, young stems and/or on fruit of P. falcatus in Ethiopian forests. Spatial analysis in the Adaba‐Dodola forest showed that disease intensity of C. uberata was significantly higher in non‐‘WAJIB’ blocks (disturbed forest) than ‘WAJIB’ blocks (sustainably managed forest) (P < 0·0001). In the temporal disease progress study, a significantly higher incidence and severity of disease on fruit was recorded during the wet season relative to dry season (P < 0·0001). The green milk stage of fruit exhibited significantly higher mean incidence (P < 0·0001) and severity (P < 0·0001) of disease compared to other growth stages of fruit. The disease incidence and severity in general, as well as on different fruit growth stages, were highly correlated (P < 0·0001, R² ≥ 0·95). Germination rate of seed decreased significantly with an increase in the level of fruit infection by C. uberata (P < 0·0001). Thus, C. uberata can apparently influence germination of seed and may pose a threat to the regeneration of P. falcatus from seeds in Ethiopian forests.