Targeted distribution of photo-assimilate in Striga hermonthica (Del.) Benth parasitic on Sorghum bicolor L.

Autotrophic photosynthesis in developing plants of Striga hermonthica, a C3 plant, parasitizing Sorghum bicolor, a C4 plant, has been quantified by measuring the ¹³C/¹²C isotope ratio (δ¹³C). This parameter has been determined in cellulose and chlorophyll extracted from plants 8 weeks after emergence. It is shown that these components are both proportionately high in autotrophic carbon in the leaf material, thus confirming the previous hypothesis that St. hermonthica photo-assimilate is directed primarily towards the synthesis of new photosynthetic capacity.     

Effect of intercropping maize and soybeans on Striga hermonthica parasitism and yield of maize

Striga hermonthica a major biotic constraint to cereal production can be controlled by trap crops. Soybean cultivars vary in ability to stimulate suicidal germination of the weed. An experiment was conducted to select soybean (Glycine max) varieties with the ability to stimulate germination of S. hermonthica seeds. Experiments were conducted with strigol Nijmegen 1® (GR 24), a synthetic stimulant, as a check. In the pot and field experiments, maize (variety WH507) was intercropped with soybeans. Variation occurred among soybean varieties in inducing germination of S. hermonthica. The relative germination induction by soybean varieties ranged from 8% to 66% compared to 70% for synthetic stimulant check. Varieties TGX1448-2E, Tgm 1576, TGX1876-4E and Tgm 1039 had the highest relative germination. Soybean varieties TGX 1831-32E, Tgm944, Tgm 1419 and Namsoy4m had high stimulation but low attachment. Intercropping maize with soybeans in the field led to a low S. hermonthica count and high maize yield.     

Characterization of nitrogen relationships between Sorghum bicolor and the root-hemiparasitic angiosperm Striga hermonthica (Del.) Benth. using K15 NO3 as isotopic tracer

The role of the host in the nitrogen nutrition of Striga hermonthica (Del.) Benth. (Scrophulariaceae) parasitic on Sorghum bicolor cv. SH4 Arval has been investigated using (15)N-nitrate as the tracer. It is shown that, when nitrate is absorbed only by the roots of the host plant, a rapid transfer of nitrogen to the parasite can be detected. The xylem sap of S. hermonthica contained approximately equal amounts of nitrate and amino acids, mostly glutamine and asparagine. Infection altered the free amino acid profile of the host tissues, leading notably to a large increase in asparagine and a decrease in glutamine. The haustoria of S. hermonthica, although rich in nitrate, showed a low concentration of free amino acids, particularly lacking in asparagine and glutamine. The roots of S. hermonthica, in contrast, were rich in both asparagine and glutamine while, in the shoots, asparagine constituted 80% of the total FAA pool. Asparagine was also found to be the primary (15)N-enriched amino acid in the shoots of S. hermonthica while, interestingly, it was glutamate that was most strongly enriched in the roots. It is concluded that nitrogen nutrition in S. hermonthica is based on a supply of both nitrate and amino acids from the host. This implies a non-specific transfer in the transpiration stream. Nitrate reduction probably occurs mainly in the leaves of the parasite. Assimilation also occurs in S. hermonthica and excess nitrogen is stored as the non-toxic nitrogen-rich compound, asparagine. This specific trait of nitrogen metabolism of the parasite is discussed in relation to the effect of nitrogen fertilization on reducing infestation.     

Gas exchange of root hemi-parasite Striga hermonthica and its host Sorghum bicolor under short-term soil water stress

The gas exchange of the upper fully expanded leaf of the root parasite Striga hermonthica and of its host Sorghum bicolor was measured under wet and dry conditions to identify the mechanisms of the devastating effects of the parasite on its hosts under drought. The short-term water stress severely reduced photosynthetic rate in infected sorghum, but less in S. hermonthica. Soil water stress did not affect leaf respiration rate in either S. hermonthica or infected sorghum. This suggests that under dry conditions both infected sorghum and S. hermonthica decreased autotrophic carbon gain. The transpiration rate of S. hermonthica, a major driving force for assimilate uptake from the host, was higher and less affected by water stress than that of infected sorghum. Stomatal density on the abaxial surfaces of the leaves was higher in S. hermonthica than in sorghum. Both S. hermonthica infection and water stress decreased stomatal conductance of the sorghum leaves. S. hermonthica, irrespective of soil water status, had greater stomatal aperture on the adaxial and abaxial surfaces of its leaves than infected sorghum. These results indicate that the higher transpiration rate of S. hermonthica even under water stress, achieved through higher stomatal density on the abaxial surfaces of the leaves and greater stomatal aperture on both surfaces of the leaves, may induce the maintenance of water and solute transfers from the host to the parasite leading to severe damage to the host under drought.     

Effect of cultural methods on Striga (Striga hermonthica (Del.)) Benth management and yield of cereals in the Savanna Zone of Nigeria: a review

Data were obtained from the research done in the Guinea Savanna (Zaria 11°11′N; 070 38′E) and Sudan Savanna (Maiduguri ?11°51′N; 13°15′E) regions of Nigeria, respectively on different cultural methods of Striga control and management. In the Guinea Savanna, trials on the effects of nitrogen on the response of resistant and susceptible upland rice varieties to Striga hermonthica infestation and the effect of resistant and susceptible varieties of maize and crop rotation on Striga infestation was carried out, while the effect of inter-cropping trap crop (Bambaranut) with resistant sorghum varieties on S. hermonthica was studied in the Sudan Savanna Zone of Nigeria. In the Guinea Savanna, it was observed that a combination of upland rice variety, Faro 40 and an application of 90 kg N/ha in the wet season and WAB 56-50 upland rice variety and 120 kg N/ha in the dry season, respectively reduced Striga infestation and produced maximum grain yield. Also, the growing of resistant variety of maize (Across 97ITZ comp. I-W) after 1 or 2 years' rotation with cowpea or soybean was observed not only to be effective in Striga control, but resulted in higher grain yield of maize. In the Sudan Savanna, the use of resistant varieties of sorghum, 1CSV1002 and 1CSV1007 intercropped with bambaranut significantly reduced Striga infestation, but the grain yield of the resistant varieties was low. From these studies, Faro 40 with 90 kg N/ha application rate and WAB56–50 with 120 kg N/ha were suitable for the management of Striga and for higher grain yield of upland rice in both wet and dry seasons, respectively, while Across 97ITZ comp. I-W, resistant maize variety and 1 or 2 years rotation with cowpea or soybean were also the best for the management of S. hermonthica and for higher maize yield in the Guinea Savanna zone. Further research needs to be carried out in the Sudan Savanna to select a high yielding resistant variety of sorghum which when intercropped with bambaranut will not only control Striga infestation but will also give high grain yield.     

Genetic Diversity of a Parasitic Weed, Striga hermonthica, on Sorghum and Pearl Millet in Mali

Eleven populations of witchweed, Striga hermonthica, were collected in four regions of Mali and investigated with 12 microsatellite markers. Extensive genetic diversity was observed, with most plants heterozygous for most markers. Allelic diversity was broadly distributed across populations with little genetic differentiation and large amounts of gene flow. Nearby fields of pearl millet and sorghum were found to have indistinguishable witchweed populations. Some population structure was apparent, but did not correlate with the local environment or host genotype, suggesting that seed transportation or other human-driven variables act to differentiate central Malian S. hermonthica populations from southern Malian populations.     

Short communication. Identification of developmentally-specific markers in germinating and haustorial stages of Striga hermonthica (Del.) Benth. seedlings

Developmentally-specific markers have been identified in the germinating and haustorial stages of Striga hermonthica seedlings. Four water-soluble proteins, preferentially expressed in germinated seedlings, were microsequenced. An haustorial-specific cDNA clone was isolated by differential screening. Tissue specificity for this clone was assessed by Northern blot hybridization analysis.Key words: Striga hermonthica (Del.) Benth, microsequencing, lipid transfer protein, superoxide dismutase.      

Impact of nitrate supply in C and N assimilation in the parasitic plant Striga hermonthica (Del.) Benth (Scrophulariaceae) and its host Sorghum bicolor L

The threshold of tolerance for nitrate of the parasitic weed Striga hermonthica (Del.) Benth and the host plant Sorghum bicolor L. was determined by estimating the impact of increasing nitrate loads on plant growth and various parameters of C and N assimilation. Nitrate supply improved chlorophyll (Chl) content and photosystem II (PSII) photochemistry of infected S. bicolor that, in comparison to S. hermonthica, displayed a low imbalance between C and N assimilation when nitrate was supplied up to 1500 mg N per plant. Indeed, nitrate supplies increased strongly the leaf N:C ratio of the parasite. The higher nitrate load induced strong accumulation of nitrate, nitrite and ammonium, and consequently the death of S. hermonthica. Nevertheless, lower nitrate loads (up to 500 mg N per S. bicolor in this study) promoted leaf expansion, PSII photochemistry and N metabolism of S. hermonthica mature (M) plants, as attested by the significant rise in soluble protein and free amino-acid contents. Following these N supplies, the nitrate tolerance of S. hermonthica was correlated with an increase in PSII activity and a high incorporation of N excess into asparagine. This confirmed the central role of asparagine in the N metabolism of S. hermonthica, although this detoxification pathway was insufficient to limit ammonium accumulation under higher nitrate loads.     

Effect of Jasmonates and Related Compounds on Seed Germination of Orobanche minor Smith and Striga hermonthica (Del.) Benth

Jasmonates and related compounds were found to elicit the seed germination of the important root parasites, clover broomrape (Orobanche minor Smith) and witchweed [Striga hermonthica (Del.) Benth]. The stimulation of seed germination by the esters was more effective than by the corresponding free acids, and methyl jasmonate (MJA) was the most active stimulant among the compounds tested.     

Effect of Growth Medium and Method of Application of Fusarium oxysporum on Infestation of Sorghum by Striga hermonthica in Burkina Faso

A 2-year (1997–1998) study was conducted at Kouaré, Burkina Faso, to investigate effect of growth medium and application method of Fusarium oxysporum isolate 4-3-B to control Striga hermonthica. In 1997, growth medium and isolate 4-3-B delayed striga emergence by 9 days. Isolate 4-3-B reduced emerged striga number by 33% compared to treatments without Fusarium. In 1998, striga emergence was delayed by 13 days by growth medium and Fusarium. The fungus reduced the number of emerged striga by 27% and, as a result, sorghum straw and grain yields were significantly improved by 10 and 38%, respectively.     

Studies on the Germination of the Seeds of Striga hermonthica: III. On the Nature of Pretreatment and after-ripening

An attempt has been made to elucidate some of the changes whichtake place in seeds of Striga hermonthica (immature, ripe andoverripe), (a) during moisture-treatment at three temperatures(15°C., 22°C., and 32°C.), and (b) during after-ripeningin dry storage. Observations on the drift of respiration of immature seed throughoutpretreatment at 15°C. show that an inverse relationshipexists between germination and respirational activity. Thissuggests that germination depends on the accumulation of somemetabolite during pretreatment. It would seem that this metabolitemay also be used as a substrate for respiration during pretreatment. After-ripening of the immature seeds has been shown to changethe nature of the seeds in some way so that their peak respirationon exposure to moisture is at first progressively reduced. Thisreduction has been ascribed to the presence of some barriercapable of restricting gaseous diffusion. The data presented, togetherwith those of the previous paperof this series, suggest that the specific metabolite requiredfor germination might be proteinaceous. Certain other data appearto rule out the suggestion that germination capacity is conditionedby the sugar content of the seeds. Reasons are given which supportthe view that one of the main effects of the stimulating solutionon the seed is to increase the permeability of the seeds togaseous diffusion.     

A study on the susceptibility of rice cultivars to Striga hermonthica and mapping of Striga tolerance quantitative trait loci in rice

Striga is a parasitic weed attacking mainly maize, sorghum, millet and cowpea. Studying the interaction between rice and Striga is valuable since rice is a model monocot. In this paper, the susceptibility of different rice cultivars to S. hermonthica was tested and quantitative trait loci (QTL) for Striga tolerance mapped on the Bala x Azucena F(6) population. Seven rice cultivars were grown with and without S. hermonthica for 14 wk. For the mapping experiment, 115 recombinant inbred lines (RILs), along with Azucena and Bala, were grown with and without Striga for 11 wk. Rice cultivars tested had different susceptibilities to Striga, ranging from highly susceptible to completely resistant. Azucena and Bala differed in the speed of Striga emergence and the impact on host growth. A genomic region between positions 139 and 166 cM on chromosome 1 was identified containing strong QTL (LOD = 4.9-15.7) for all traits measured. This indicates that genes for Striga tolerance exist in rice germplasm and the mapped QTL can be further studied to promote understanding of the nature of resistance/tolerance and breeding for Striga-resistant crop plants.     

Studies on the Germination of the Seeds of Striga hermonthica: II. THE EFFECT OF THE STIMULATING SOLUTION ON SEED RESPIRATION

1. Data are presented which show the effect of the natural stimulatingsolution on the respiration of the seeds of Striga hermonthica.
2. The stimulating solution has been shown to enhance the aerobicrespiration of seeds exposed to it, compared with that of seedsto distilled water at the same temperature. This effect on therespiration of the seeds is quite independent of any previousmoisture-treatment of the seeds, and thus of germination.
3. Themaximum anaerobic rate of carbon-dioxide output of air-dryseedsafter treatment with the stimulating solution may be upto fifteentimes greater than that of seeds moistened with distilledwater.The anaerobic respiration of air-dry seeds after exposuretothe stimulating solution is greatly reduced by 0?025 M. sodiumfluoride and 0?001 M. sodium monoiodoacetate.
4. The stimulatingsolution has been found to have little effecton the anaerobicoutput of carbon dioxide from seeds which hadbeen moisture-treatedfor 6 days at 22? C. before treatmentwith the stimulating solution.
5. No correlation could be established between the effects ofthestimulating solution on germination and respiration.

     

The parasitic angiosperm Striga hermonthica can reduce photosynthesis of its sorghum and maize hosts in the field

Two cultivars of sorghum (CK60 and Ochuti) and one cultivarof maize (H511) were grown in field plots in western Kenya inthe presence or absence of the parasitic angiosperm Striga hermonthica,with or without a single addition of nitrogen fertilizer (150kg N ha^–1) using a factorial design. A progressive declinein rates of photosynthesis of Striga-infected plants were observedfor the sorghum cultivar CK60 from 30 d after planting (DAP)and for maize from 40 DAP, until measurements ended 63 DAP.At this time photosynthetic rates were 46% and 31% lower inthe Striga-infected sorghum and maize cultivars, respectively,compared to uninfected control plants. No decline in photosynthesiswas observed in the second sorghum cultivar studied, Ochuti,a local land race reported to show some tolerance to the parasite.The trends in photosynthesis reflected stunting of the cereals,as determined by the height of the youngest emerged ligule,however, only the grain yield of the sorghum cultivar CK60 wassignificantly reduced by the presence of the parasite. The nitrogenapplication influenced neither the growth nor the photosyntheticparameters measured, and possible explanations for the absenceof responses are discussed. It is concluded that S. hermonthicacan reduce photosynthetic rates of field-grown sorghum and maize,and suggest that an ability to maintain high rates of photosynthesiswhilst infected may be an important correlate of tolerance tothe parasite. Key words: Parasitic angiosperm, photosynthesis, nitrogen, tropical weeds, tropical agriculture     

The effect of light on the Og of plant tissues. I

Summary The increment of O_g in the guard cells in response to light as reported by previous investigators was found to be true qualitatively but not quantitatively.A special device was constructed, which permitted the elimination of the heat factor when the leaves were irradiated.In experiments 1 and 2 the relative values of the bell and the device were determined.In experiments 3 and 4 the relative effects of light were studied on younger and older leaves.In experiments 5 and 6 the effects were studied on old and young leaves ofHedera helix. This plant serves as a standard for comparisons of results obtained by subjecting plants to various factors of the environment.In experiments 7 and 8 the effect of light on young, middle-aged and old leaves ofHedera helix, when water was supplied, in one case but not in the other were studied. In experiment 8 the same leaf was treated in the device and then in the dark chamber, and once more in the light. The results are particularly enlightening.In the supplementary experiments, it was verified that the phenomenon of an increment of the guard cells and a decrement of the spongy parenchyma in O_g, when irradiated, is probably general.     

The use of polyethylene bags in the culture and study of Striga spp. and other organisms on crop roots

A technique is described by which root parasites such as Striga and Orobanche species may be grown on the roots of suitable host plants growing on a layer of glass-fibre filter paper in a flattened, standard polyethylene bag. The single-bag units are compact, inexpensive, simple to set up and allow repeated observation without damage. Some other potential uses of the technique are described.     

The strigolactone germination stimulants of the plant-parasitic Striga and Orobanche spp. are derived from the carotenoid pathway

The seeds of parasitic plants of the genera Striga and Orobanche will only germinate after induction by a chemical signal exuded from the roots of their host. Up to now, several of these germination stimulants have been isolated and identified in the root exudates of a series of host plants of both Orobanche and Striga spp. In most cases, the compounds were shown to be isoprenoid and belong to one chemical class, collectively called the strigolactones, and suggested by many authors to be sesquiterpene lactones. However, this classification was never proven; hence, the biosynthetic pathways of the germination stimulants are unknown. We have used carotenoid mutants of maize (Zea mays) and inhibitors of isoprenoid pathways on maize, cowpea (Vigna unguiculata), and sorghum (Sorghum bicolor) and assessed the effects on the root exudate-induced germination of Striga hermonthica and Orobanche crenata. Here, we show that for these three host and two parasitic plant species, the strigolactone germination stimulants are derived from the carotenoid pathway. Furthermore, we hypothesize how the germination stimulants are formed. We also discuss this finding as an explanation for some phenomena that have been observed for the host-parasitic plant interaction, such as the effect of mycorrhiza on S. hermonthica infestation.