Phelipanche ramosa is a major weed holoparasite characterized by a broad host range with a suboptimal development on numerous hosts, suggesting inter- or intra-species specificities. Seeds of P. ramosa germinate after exposure to exogenous chemicals exuded by surrounding host roots such as strigolactones, the concentrations of these germination stimulants varying between hosts. In France, P. ramosa is characterized by genetically differentiated populations presenting varying germination rates and a host specificity. The objective of our study was to investigate the sensitivity of seeds of two P. ramosa populations harvested on tobacco and oilseed rape, to a set of GR24 concentrations, a synthetic strigol analogue. The assessment of the germination rate was based on in vitro experiments. Seeds of P. ramosa were placed in Petri dishes with various concentrations of GR24. The cumulative number of germinated seeds of P. ramosa was counted several times after application of the treatment. Cumulative germination curves were analysed using a three-parameter log-logistic model and a time-to-event approach. The results show that the germination rate of P. ramosa seeds depends on the GR24 concentration and the duration of stimulation, but also that the response to these two factors varies greatly according to the origin of the P. ramosa seeds. The difference in germination speed between P. ramosa populations further shows distinct responses at the intraspecific level, thus suggesting that the specialization of P. ramosa probably occurs at least from the first stage of the holoparasite cycle.

Host-specific interactions can maintain genetic and phenotypic diversity in parasites that attack multiple host species. Host diversity, in turn, may promote parasite diversity by selection for genetic divergence or plastic responses to host type. The parasitic weed purple witchweed [Striga hermonthica (Delile) Benth.] causes devastating crop losses in sub-Saharan Africa and is capable of infesting a wide range of grass hosts. Despite some evidence for host adaptation and host-by-Striga genotype interactions, little is known about intraspecific Striga genomic diversity. Here we present a study of transcriptomic diversity in populations of S. hermonthica growing on different hosts (maize [Zea mays L.] vs. grain sorghum [Sorghum bicolor (L.) Moench]). We examined gene expression variation and differences in allelic frequency in expressed genes of aboveground tissues from populations in western Nigeria parasitizing each host. Despite low levels of host-based genome-wide differentiation, we identified a set of parasite transcripts specifically associated with each host. Parasite genes in several different functional categories implicated as important in host–parasite interactions differed in expression level and allele on different hosts, including genes involved in nutrient transport, defense and pathogenesis, and plant hormone response. Overall, we provide a set of candidate transcripts that demonstrate host-specific interactions in vegetative tissues of the emerged parasite S. hermonthica. Our study shows how signals of host-specific processes can be detected aboveground, expanding the focus of host–parasite interactions beyond the haustorial connection.

Selective control of Egyptian broomrape is extremely difficult because the close association between host crop and parasite limits the use of most mechanical and herbicidal approaches. However, this host–parasite interaction can also form the basis of the simplest control strategy: parasite-resistant crops. Although much work has been conducted to identify and characterize mechanisms of parasite resistance, varieties with stable resistance are still unavailable for most affected crops. The development of resistant crops can be accelerated by genetic engineering to the extent that important aspects of the host–parasite interaction are understood. In this study, we characterize a variety of gene promoter elements with respect to parasite induction and expression pattern. Studies were conducted using transgenic plants expressing fusions of the β-glucuronidase reporter gene with promoter elements from several genes. Promoters from genes known to have increased expression in response to pathogen attack or wounding showed localized, induced expression after parasitism. These included phenylalanine ammonia lyase, chalcone synthase, sesquiterpene cyclase, and HMG1 (3-hydroxy-3-methylglutaryl CoA reductase). In contrast, the systemic acquired resistance–associated gene PR-1a was not induced by parasitism. Non–defense-related genes varied in response, with squalene synthase being repressed, whereas farnesyltransferase was highly expressed in the region of parasite attachment. These results demonstrate a range of expression, both in intensity and tissue specificity, in response to parasitism.

Broomrapes (Orobanche spp.) are chlorophyll-lacking root parasites of many dicotyledonous species and cause severe damage to vegetable and field crops from several botanic families such as Fabaceae, Solanaceae, Compositae, and Umbelliferae. In Oregon, small broomrape has been identified as a parasite of red clover. In Oregon field studies, small broomrape control was excellent when imazamox was applied postemergence to red clover but preemergence to small broomrape. Temperature is one of the main factors that affect broomrape development. The objective of this study was to optimize small broomrape chemical control in red clover based on growing degree days (GDD). The study was conducted in controlled temperature conditions. Red clover plants were grown in soil artificially infested with small broomrape seeds. Imazamox was applied at 800, 1,000, 1,200, and 1,400 GDD. There was no injury to red clover from any imazamox treatment at any of the application timings. Small broomrape shoot emergence was reduced where imazamox was applied compared to the untreated control. Early imazamox applications reduced small broomrape biomass more than later applications. Control was greatest when imazamox was applied at 20 g ai ha−1 at 1,000 GDD. This application controlled small broomrape for 800 GDD after initial treatment. However, season-long control would require an additional treatment. This model predicts the optimal timing and rate of imazamox application for small broomrape control in red clover.

Small broomrape is a holoparasitic plant that attaches to the roots of red clover as well as several other host plants. Hosts and false hosts produce stimulants that induce small broomrape germination but small broomrape does not attach to a false host. Wheat has been identified as a false host for small broomrape; therefore, studies were conducted to investigate the effect of red clover and wheat root exudates on small broomrape germination. In one study, the effect of exudates from red clover and wheat at multiple growth stages on small broomrape germination was evaluated. Red clover induced small broomrape germination at all growth stages tested but was greatest (78%) in the presence of exudates from red clover at the three-trifoliolate stage. Maximum small broomrape germination was 25% when exposed to exudates produced by one-leaf-stage wheat. In a second study, the relationship between small broomrape germination and host growth condition was evaluated using root exudates from red clover or wheat grown under several temperature conditions for either 4 or 8 wk. For the different temperatures, there were no differences in small broomrape germination when exudates of red clover grown for 4 wk were used. Small broomrape germination was reduced when exposed to exudates from red clover plants grown for 8 wk at 10 C compared with plants grown at 15, 20, and 25 C. Differences in small broomrape seed germination were observed with temperature under which wheat was grown for 4 wk, but not for 8 wk. Although wheat exudates resulted in less small broomrape seed germination than red clover exudates, growing wheat as a false host in a small broomrape-infested field could be an important component of an integrated management plan.

Small broomrape is an annual holoparasitic weed that was recently discovered in red clover production fields in Oregon. Imidazolinone herbicides such as imazamox control small broomrape; however, the mechanism of uptake by the parasite is largely unknown. Studies were conducted to determine the imazamox route of uptake by small broomrape in red clover, and to determine the potential for imazamox to be exuded from red clover and the subsequent effect on small broomrape. Small broomrape control was best at 90% when imazamox was foliar-applied, and worst at 42% or less when imazamox was soil-applied. The presence of activated charcoal to adsorb imazamox at the soil surface did not affect efficacy of broadcast foliar treatment. Small broomrape control was also evaluated when a foliar-treated red clover plant was grown in the same pot as a nontreated, parasitized red clover plant that was bagged during herbicide application. Activated charcoal was spread on the soil surface to adsorb imazamox, thus limiting herbicide uptake routes to the foliage of one of two red clover plants in the pot. Small broomrape attachment decreased on nontreated red clover when the other red clover plant in the pot was treated, suggesting roots exuded the herbicide or an active metabolite.

Small broomrape is a parasite of several broadleaf plant species. Consequences of small broomrape infestation in host cropping systems include seed contamination, reduction in crop seed yield, and host plant death. The effect of small broomrape parasitism on the biomass partitioning of its primary host, red clover, has not been documented. Greenhouse experiments were conducted to determine the relationship between small broomrape and red clover biomass accumulation. Total biomass of parasitized red clover plants was 15 to 51% less than nonparasitized red clover plants. Small broomrape parasitism reduced the amount of dry matter allocated to red clover inflorescences by 50 to 80%. Small broomrape dry matter accumulation was strongly related to total red clover–small broomrape dry matter accumulation. Small broomrape attachment number per red clover plant was a poor indicator of relative small broomrape dry weight accumulation. The results of this study indicated that small broomrape accumulated resources from red clover at the greatest expense to the economically important reproductive tissues.

The influence of the holoparasite branched broomrape on the vegetative growth, leaf chlorophyll content, photosynthetic rate, and chlorophyll fluorescence of tomato was studied over two growing seasons on plants grown in a commercial greenhouse. The presence of the parasite strongly reduced the aerial biomass by acting as a competing sink for assimilate, but more importantly, by compromising the efficiency of carbon assimilation via a reduction in leaf chlorophyll content and photosynthetic rate. The chlorophyll fluorescence parameters F0, Fm, Fv, and Fv/Fm were all altered in parasitized plants, indicating that branched broomrape–infected plants are more susceptible to photoinhibition. The degree of damage to the host was not dependent on either the number or the biomass of parasitic plants per host plant. We suggest that the ability to maintain a high photosynthetic rate, leaf chlorophyll content, or both and the ability to minimize photoinhibition can be developed as indirect assays for improved tolerance to branched broomrape.

Small broomrape is a federally listed noxious weed that was first identified in Oregon in 1923. Between 1923 and 1997, there were only six reports of the species in Oregon. Small broomrape is a holoparasitic weed that attaches to the root of its host and can lead to crop failure depending on the level of infestation. Small broomrape was identified in a red clover field in Oregon in 1998. In 2000 and 2001, the Oregon Department of Agriculture surveyed red and white clover fields in Oregon for the presence of small broomrape; 15 infested fields were identified in 2000 and 22 in 2001. The source of the small broomrape seed was not identified although there was speculation that small broomrape seed may have been contaminated clover seed stock. In 2000, a quarantine was established for small broomrape in red clover seed. Research identified effective, easy to implement management strategies for small broomrape including the use of false host crops and imazamox. In 2003, the ODA amended the quarantine to eliminate the mandatory seed sampling and testing and the requirement of reporting infested fields and seed lots. In 2011, there are still populations of small broomrape present in red clover fields in Oregon.

The facultative hemiparasite Rhinanthus minor was grown alone, or with either Hordeum vulgare or Trifolium alpestre as a host. All plants were fed 5 mM nitrate. In the parasites, successful attachment led to dramatic increases in growth (particularly with barley as host) and in tissue concentrations of total N, NO3−, H2PO4− and K+, as well as moderate increases in Cl− and SO42−. Mg2+ and Ca2+ concentrations either remained unchanged, increased or decreased, depending on the host. Xylem sap was collected from leaf veins (barley) or the stem (Rhinanthus and clover), by applying pneumatic pressure to the rooted soil to raise the soil water potential so that in vivo xylem sap exuded from small incisions once the applied pressure balanced the tension exerted by transpiration. High balancing pressures were needed for unattached Rhinanthus, whereas simultaneous collection of xylem sap from attached Rhinanthus and its host, at a much lower balancing pressure, was possible only when the parasite was enclosed in polyethylene film and high transpiration abolished. From plots of xylem sap volume flow versus applied pressure, the hydraulic conductivity Lp and the overall hydraulic resistance r were obtained. The value of r was high in unattached Rhinanthus and dramatically decreased by attachment to a host, highlighting the improved access to water gained by Rhinanthus when parasitizing a host. For the parasites, attachment resulted in greatly increased concentrations and solute flow rates in xylem of K+, NO3−, H2PO4− and amino acids, and decreased concentrations of Ca2+ and Mg2+. Estimates of NO3− reduction in the parasite showed that 99% was reduced in unattached plants, 85% in those attached to clover and 52% in those attached to barley. In the parasitized hosts, ion concentrations in xylem sap were somewhat increased relative to unparasitized controls. Conversely, xylem sap amino-acid concentrations in infected hosts were decreased. Glutamine (Gln) was the principal amino acid in xylem sap of unattached Rhinanthus, but after attachment to both hosts asparagine (Asn) predominated. In clover, Asn was the major transport amino acid, suggesting massive transfer of Asn from host to parasite. In unparasitized barley, however, Gln was the major xylem sap amino acid, but unexpectedly, in this species Asn was induced as the principal amino acid when parasitized. The results are discussed in terms of mutual host–parasite interactions.