Reduction of Striga hermonthica parasitism on maize using soybean rotation

Striga hermonthica is a serious parasite of cereals in most of the semi-arid savanna zone of West Africa, causing substantial yield loss. It has been observed that some soybean cultivars are capable of stimulating germination of S. hermonthica seed, which would reduce the seed bank in the soil. This study was undertaken to quantify the effect of a soybean crop (compared with a sorghum control) on S. hermonthica emergence in subsequent maize in three farmers' fields in northern Nigeria. Soybean cultivar TGx 1740-7F, previously identified as efficacious for S. hermonthica seed germination, was grown at four densities without P fertilizer to test the effect of increasing plant density of soybean on subsequent S. hermonthica parasitism. The effect of P as single super phosphate was tested on the two highest soybean densities. S. hermonthica parasitism on maize was significantly lower after unfertilized soybean than after the sorghum control treatment at two of three trial sites. Soybean rotation increased maize yield by approximately 90% for the three sites combined. Increasing soybean plant density did not result in lower emerged S. hermonthica. Application of P to soybean at the higher soybean densities resulted in higher root length density, lower emerged S. hermonthica on maize (P<0.15), and significantly higher maize yield. The results suggest that an efficacious cultivar of soybean reduces S. hermonthica parasitism on a succeeding maize crop and that the effect is increased by application of P to the soybean.     

Wheat (Triticum aestivum) Is Susceptible to the Parasitic Angiosperm Striga hermonthica, a Major Cereal Pathogen in Africa

ABSTRACT Striga hermonthica is a parasitic weed endemic to sub-Saharan Africa. It most commonly parasitizes sorghum, maize, pearl millet, and upland rice, lowering yields and affecting the welfare of over 100 million people, principally subsistence farmers. Cereal crops with complete resistance to this pathogen have not been reported. In southern and eastern Africa, where Striga spp. are endemic, 5.6 million ha of wheat are cultivated annually. Despite this, there are only isolated field reports of wheat infected with Striga spp. It is not clear whether this is due to resistance in this cereal or to environmental factors. In this article, we examined the ability of root exudates from five cultivars of wheat (Chablis, Cadenza, Hereward, Riband, and Brigadier) to trigger germination of S. hermonthica seed. A study of the development of S. hermonthica on two cultivars of wheat (Hereward and Chablis) and on a range of ancestral relatives of wheat (Triticum and Aegilops spp.) then was conducted. Last, the effect of Striga spp. on host growth and yield was examined using wheat cv. Chablis and compared with that of a highly susceptible sorghum cultivar (CSH-1). Wheat was able to support the germination, attachment, and subsequent development of Striga spp. All wheat cultivars and ancestral species of modern wheat (Triticum and Aegilops spp.) were susceptible to S. hermonthica. In addition, in wheat, infection severely lowered plant height (-24%) and biomass accumulation (-33%); a small parasite biomass elicited a large host response. In conclusion, wheat is highly susceptible to S. hermonthica and, in light of global climate change, this may have implications for wheat-producing areas of Africa.     

Desmodium species and associated biochemical traits for controlling Striga species: present and future prospects

We discovered serendipitously a new and highly effective intervention against Striga spp., including Striga hermonthica (African witchweed), in cereals, which involves intercropping with cattle fodder legumes, Desmodium spp., including D. uncinatum . Although soil shading and additional nitrogen made some contribution to the reduction of S. hermonthica infestation, an allelopathic mechanism associated with the intercrop was a major factor. Root exudates of D. uncinatum contain novel flavonoid compounds, some of which stimulate germination of S. hermonthica and others dramatically inhibit its subsequent development, including radicle growth. Desmodium spp. have been developed as intercrops for both maize and sorghum and are now being evaluated for millet. From the experience with Desmodium spp., there is now the possibility for producing edible legumes suitable for intercropping with maize and other cereals to respond to a broader profile of farmer practices. These legumes would incorporate the powerful S. hermonthica controlling properties of D. uncinatum through feasible breeding programmes, with appropriate contributions from analytical chemistry to plant molecular genetics and biotechnology. In the longer term, it may also be possible to transfer genes associated with the allelopathic attributes to cereals themselves by heterologous gene expression, creating a new generation of parasitic weed-free cereals.     

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.     

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.     

Can wild relatives of sorghum provide new sources of resistance or tolerance against Striga species?

Summary The genus Striga contains some of the most noxious parasitic plants, which have a devastating impact on cereal production in Africa; of most importance are Striga hermonthica and Striga asiatica . Complete resistance to infection by Striga species does not exist in cultivated cereals. Of great interest is the possibility that wild relatives of cereals may provide a genetic basis for resistance or tolerance to infection and may be of enormous value for the development of resistant crops. A wild relative of cultivated sorghum, Sorghum arundinaceum , demonstrated tolerance to infection by S. asiatica , with little impact of S. asiatica on host growth or grain production compared with the detrimental impact of the parasite on cultivated sorghum. Infection by S.hermonthica , however, had a significant influence on host performance for both wild and cultivated sorghum. Differences in host:parasite responses may be explained by the timing of parasite attachment and differences in host specificity for these two Striga species.     

A study of assimilation and translocation in Cuscuta hyalina Heyne ex Roth., Orobanche ramosa L. and Striga hermonthica Benth.

Quantitative determinations of chlorophyll showed that Cuscuta hyalina and Striga hermonthica possess chlorophyll. No trace of chlorophyll was found in Orobanche ramosa. Autoradiographic evidence showed that both radiocarbon and radiophosphorous moved from host to parasite in alt three species. Photosynthesis occurs in S. hermonthica when the parasite is isolated from its host. The adventitious roots of S. hermonthica and O. ramosa appear to be functional in absorbing a proportion of the required inorganic substances and water. S. hermonthica is a hemiparasite, O. ramosa is a holoparasite but C. hyalina is best described as a partial parasite.     

Weed flora and soil seedbanks in fields dominated by Imperata cylindrica in the moist savannah of West Africa

Imperata cylindrica (L.) Raeuschel is a dominant and infamous grass weed in the savannah of West Africa. Research to reduce the weed to non-damaging levels is a priority activity at many agricultural institutions. The successful development and implementation of long-term I. cylindrica management strategies depend on the ability to predict changes in weed composition after I. cylindrica has been controlled effectively. The weed flora and soil seedbank were assessed from 329 fields dominated by this species in the fringes of the humid forest (HFF), coastal/derived savannah (CDS) and in the southern Guinea savannah (SGS) in 1996 and 1997. The objectives of the study were to correlate species composition of the weed flora with that of the soil seedbank and to determine the effect of management factors and soil properties on the composition of the weed flora. Species richness in the weed flora and in the weed seedbank was higher in the SGS than in the CDS and HFF. Mean weed density per field was generally higher in the HFF (156 ± 25.0 weeds m^–2) than in the CDS (108 ± 8.1 weeds m^–2) and in the SGS (92 ± 6.3 weeds m^–2). Weed composition varied with agroecological zone as well as with management factors and soil properties. Sørenson's index of similarity was low (mean=0.20) in all zones, indicating poor similarity between the weed flora above-ground and the soil seedbank.     

Assessment of management options for Salsola australis in south-west Australia by transition matrix modelling

A matrix model of the life cycle of Salsola australis was constructed, based on population ecology data collected from the district of Lake Grace, Western Australia. The model was used to assess potential control strategies for this summer annual weed within the Western Australian broad acre grain cropping system. The population growth rate (λ) of S. australis in the absence of weed control strategies was 1.49 and was virtually unaffected by the dormant seedbank. However, λ increased to 8.21 if it was assumed that a constant number of seed immigrated into the area in question from neighbouring populations of S. australis , through farm-scale seed dispersal. As a result, effective weed management depended on reducing seed dispersal. The model determined that burning all senesced, mobile plants in late autumn, combined with herbicide control of the largest cohorts of S. australis in summer and autumn, reduced population growth rate to 0.79. This control strategy resulted in a 66.1% chance of the population becoming extinct over 25 years. Management strategies are proposed based on the results of the models and further research is required to validate their effectiveness and practicality in the field.     

Impact of cropping systems on the weed seed banks in the northern Great Plains, USA

Very little research has been done on weed seedbank communities in the northern Great Plains region of the USA. The objectives of this study were to evaluate the effects of management systems on the weed seed banks and to assess the relationship between the weed seed banks and the above-ground weed communities. The seed banks were sampled along a range of areas of above-ground weed diversity in organic and conventional spring wheat production fields near Big Sandy, Montana, over 2 years. Eight 1 m × 0.33 m areas were selected in each field to encompass a wide range of above-ground weed diversity and soil cores were taken in each area to a depth of 20 cm and split in half at 10 cm. The number of seeds recovered from the top 10 cm of soil and the 10–20 cm depth were significantly affected by the sampling year and an interaction between the cropping system and the sampling year. The year was the only significant predictor affecting the weed seedbank diversity at both depths. A significant interaction between the management system and the year observed at the 10–20 cm sampling depth reflected the higher rate of decrease in diversity occurring between 2005 and 2006 in the conventionally managed fields compared with the organic fields. A multivariate ordination indicated that, although the year played a significant role in determining the weed seedbank communities, the management system had a role only during 2006. We failed to detect strong correlations between the above-ground and underground weed communities. The results of this study suggest that, in the studied region, yearly fluctuations in environmental factors have significant impacts on the weed seed banks.     

A comparative study on Striga hermonthica interaction with a sensitive and a tolerant sorghum cultivar

In this study, the effects of Striga hermonthica (Del.) Benth. on a sensitive and a tolerant sorghum genotype were investigated, and the underlying tolerance mechanisms were distinguished. The sensitive sorghum cv. CK60-B and the tolerant sorghum landrace Tiemarifing were grown in pots with and without seed infestation. Both sorghum genotypes responded to infection by the parasite, but it was evident that CK60-B was more strongly affected than Tiemarifing. Sorghum plant height, final leaf number, green leaf area, kernel yield, number of kernels and 1000-kernel weight were significantly reduced by infection, which also had a marked effect on the phenological development of CK60-B; the majority of the plants remained vegetative and, in the remaining plants, flowering was delayed by about 2 weeks. No effect on the phenological development of Tiemarifing was observed. The tolerant landrace showed significantly lower and delayed emergence of S. hermonthica than the sensitive cultivar, and this could be explained by a delay in the onset of attachment. Significantly higher numbers of reproductive parasitic plants were found in the pots with the sensitive sorghum plants. It is concluded that differences in root architecture and the resulting early infection and higher S. hermonthica numbers are partly responsible for the stronger effects of the parasite on the sensitive cultivar.     

Effects of soil and crop management practices and pedo‐hydrological conditions on the seedbank size of Galinsoga spp. in organic vegetable fields

Galinsoga quadriradiata and Galinsoga parviflora are very troublesome weeds in many organic vegetable crops in Europe. A very straightforward method to keep Galinsoga infestations under control is by targeting the Galinsoga seedbank. To identify cropping systems able to reduce the seedbank size in vegetable‐based cropping systems, the relationships between the seedbank size of Galinsoga species and prevailing soil/crop management practices and pedo‐hydrological conditions were investigated. Hereto, the seedbank of the 0–20 cm topsoil layer was sampled in 50 organic vegetable fields and analysed according to the seedling emergence method. Field history data were collected for the past 5 years, and physical, chemical and microbial soil quality was determined. Galinsoga quadriradiata was the most frequent and abundant Galinsoga species in the weed seedbank. The genus Galinsoga was present in 90% of the soil weed seedbanks of organic vegetable fields but displayed wide variation in abundance. Smallest Galinsoga seedbanks were found in fields that were predominantly tilled with non‐inversion implements or rotationally ploughed, and continuously cropped with competitive crops during the entire growing season (April 15‐November 15). Contrary to G. quadriradiata, seedbank size of G. parviflora was closely related to soil organic carbon content and sand fraction. Remarkably, soils with a low level of easily plant‐available phosphorus and concomitant high activity of arbuscular mycorrhizae had smaller G. quadriradiata seedbanks. To reduce Galinsoga infestations, fields should preferably be tilled without soil inversion, fertilised with organic amendments with low content of readily plant‐available phosphorus and cropped with competitive crops all season long.     

Evaluation of variability in Striga aspera, Striga hermonthica and their hybrids using morphological characters and random amplified polymorphic DNA markers

Striga aspera and Striga hermonthica are recognized as separate species, but their close morphological similarity causes difficulty in distinguishing between them in areas where they coexist in Africa. In this study, crosses between the species were made using randomly selected morphologically typical parental plants collected from different locations in Nigeria. Genetic analysis of both species and their reciprocal F₁ hybrids were determined using cluster analysis of DNA profiles derived from genetic polymorphism (RAPD)-polymerase chain reaction (PCR) markers. Principal component and hierarchical cluster analyses were used to separate parental and hybrid populations based on 13 morphological characteristics. Morphological data from wild samples of both species were compared with the hand-pollinated parental, F₁ and F₂ hybrids, and back-crosses. Results showed that S. aspera and S. hermonthica were genetically and morphologically distinct. Morphological and genetic analyses revealed two major clusters: a S. aspera cluster and a S. hermonthica cluster. Genetically, the F₁ hybrids showed closer affinity to their maternal parents, while morphologically, the F₁ hybrids formed distinct clusters intermediate to the parents. Most F₂ plants and back-crosses were morphologically similar to S. hermonthica . Comparative morphological analysis of wild and hand-pollinated populations showed some samples from the wild clustered with the hybrids, suggesting that hybrids may exist in nature.     

Reproductive Ability of Hybrids of Striga aspera and Striga hermonthica

ABSTRACT Striga aspera and S. hermonthica are sympatric in Africa. Each may serve as virulent gene reservoirs for the other if they hybridize and their hybrids are virulent and fertile. Intraspecific and interspecific crosses were made within and between the species, and reproductive success was determined. Freshly harvested seeds from the parental and F(1) crosses were tested over time for germination. Chromosome counts from shoot-tip squashes of seedlings of S. aspera were determined as n = 18, and as n = 19 for S. hermonthica. Hybridization results indicated that S. aspera and S. hermonthica could be intercrossed and their hybrids successfully backcrossed to either parent. Reproductive success in all crosses ranged from 68 to 95% and seeds of all the crosses were viable, germinated on exposure to a synthetic germination stimulant, and were pathogenic on maize. Seeds from the hybrids and backcrosses were less viable and germinable than either parent, suggesting that the two species were not 100% compatible. Results suggest that the two species are closely related, but are separate taxa, perhaps at the subspecies level. Seed dormancy for both species and the F(1) hybrids was less than 84 days after pollination. Germination of S. hermonthica seeds reached 31% at 28 days after pollination. Frequently occurring seed germination peaks were observed for all the seeds tested.     

Modelling population dynamics of Sinapis arvensis in organically grown spring wheat production systems

Although we know that alterations in crop density, crop spatial pattern and inclusion of more selective weed control can improve weed suppression for organic growers, it is unknown whether these result in changes to the weed seedbank that increase cropping system profitability over time. Data collected from field trials conducted in 2009 and 2010 in Maine, USA, comparing regional grower practices (Standard) with management that aimed to (i) facilitate better physical weed control through the use of wide row spacing and inter‐row cultivation (Wide) or (ii) enhance crop–weed competition through increased seeding rate and narrow inter‐row spacing (Narrow HD), were used to construct a matrix population model with an economic sub‐model. Using field measurements of grain yield and weed survival and fecundity, we investigated the lasting implications of employing alternative organic spring wheat (Triticum aestivum) production practices on Sinapis arvensis population dynamics. In most scenarios, the model indicated that regional production practices were not sufficient to prevent an increase in the weed seedbank, even with excellent weed control. The two alternative methods, on the other hand, were able to limit weed population growth when initial densities were low or cultivation efficacy was >80%. Due to higher seed costs in the Narrow HD system, net returns were still lower after 10 years of simulation in this system compared with wide rows with cultivation, despite a lower weed seedbank.     

A note on the chemical composition, intake and digestion of Striga hermonthica herbage by sheep

Striga hermonthica causes serious crop yield losses in West Africa. Hand pulling, an effective method for the reduction of light infestations, might be encouraged if farmers could use this weed as livestock feed. This study evaluated the chemical composition and the voluntary intake and digestion of S. hermonthica herbage by sheep. Crude protein (g kg⁻¹ dry matter (d.m.)) was 184 in the whole plant, 230 in the leaf and 87 in the stem. Ash content varied from 183 to 253 g kg d.m.⁻¹. The concentration of neutral and acid detergent fibre and lignin in whole pot-grown plants was 364, 278 and 127 g kg d.m.⁻¹ respectively. The digestibility of dry and organic (o.m.) matter was 493 and 657 g kg⁻¹, respectively, and intake of digestible o.m. was 27.1 g kg W^−0.75. The relatively high N and P levels in S. hermonthica warrant further evaluation in terms of its potential use as a source of protein or for compost. Its use as a feed appears to be limited by the high ash content and possibly by anti-nutritional effects on animals. These effects should be further investigated before recommending its use for this purpose.     

Biology of Striga hermonthica (Scrophulariaceae) in Sahelian Mali: effects on pearl millet yield and prospects of control

Aspects of the population: dynamics of Striga hermonthica (Del.) Benth. on pearl millet (Pennisetum americanum (L.) K. Schum.) were studied in Sahelian Mali. Seasonal development was recorded and density-dependent mortality and fecundity in post-emergent populations investigated. An attempt was also made to assess yield loss due to S. hermonthica attack using a regression approach. The first S. hermonthica plants emerged 42 days after crop germination, and the minimum time to complete the life cycle from emergence was 56 days. Results indicated a premature mortality of 66% in emerged populations. The growth stage reached by the time of host plant death appeared to be important in determining survival to maturity, premature mortality being greatest in plants that had not reached flowering by this time. The potential seed output per plant was estimated at 10 985. There was no evidence that the fecundity of S. hermonthica was affected by the density of emerged populations. Regression models showed a significant relationship between millet yield and emerged S. hermonthica density. The results suggest that lower densities give a relatively larger decrease in yield per S. hermonthica plant. The results of the study are discussed in relation to their implications for control.     

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.     

Reduction in field emergence and seedbank density of Galinsoga quadriradiata and other weeds after contrasting false seedbed strategies in organic vegetable fields

Mechanical weed control in low competitive, organic vegetable production systems is challenging, particularly in fields with large populations of Galinsoga spp. (Asteraceae). Various false seedbed techniques are used prior to crop planting or sowing to prevent weed emergence, albeit with variable success. This study investigated the impact of machinery type (flamer, hoe and harrow), number of passes (2 and 4), tillage depth (1–4 cm) and intensity (double and single hoeing, and hoeing with or without additional harrowing) on weed emergence and seedbank density in 0–5 cm topsoil of organic vegetable fields. False seedbed machinery that did not or minimally disturb the soil was most appropriate for preventive control of Galinsoga quadriradiata (Hairy galinsoga) and total weed seeds, with reductions in seedling emergence up to 99% and 73%, respectively, for flaming, and 74% and 67%, respectively, for 1 cm deep hoeing, 1 month after false seedbed creation. Compared with 1 cm deep hoeing, 1 cm deep harrowing was 16% less effective in the control of emerged seedlings, while flaming was highly effective in preventing weed seedling emergence, even after a low number of passes. Tillage intensity was less important than tillage depth for the reduction in weed emergence and seedbank density. Overall, tillage was more effective for seedbank reduction than flaming.     

Soil seedbanks in slash‐and‐burn rice fields of northern Laos

Crops in shifting cultivation fields often suffer from severe weed infestation when long fallow periods are replaced by short fallow periods. The soil seedbank as a source of weed infestation was studied in four fields that differed in their last fallow duration. The effect of burning was analysed by comparing adjacent pre‐burn and post‐burn samples (two sites). Surface vegetation was monitored from burning to harvest in the plots from which soil samples were taken to determine the fraction of the seedbank germinating (three sites). Seedbank size (1700–4000 seedlings m^?2) varied depending on a single species, Mimosa diplotricha. Burning reduced emergence of most species, but stimulated emergence in others. Densities in the seedbank were not correlated with above‐ground abundances in the field, except for some species. Most species emerging after 50 days from the soil samples (40% of seedlings) were absent from the field after 190 days. Whilst the data from this study are derived from only four fields, the weed problems after short‐term fallowing appeared to be due to a larger fraction of the seedbank emerging, possibly due to shallow burial, and to a floristic shift towards adaption to burning, rather than the size of the seedbank per se.