Gene flow between alien and native races of the holoparasitic angiosperm Orobanche minor (Orobanchaceae)

The holoparasitic angiosperm Orobanche minor parasitizes a diverse range of flowering plants from at least 16 orders in both the monocots and eudicots. However, populations of O. minor show host specificity at a local level. Our previous work identified the potential for host specificity to act as a catalyst for genetic divergence among populations of O. minor. Here we have extended this investigation by sampling populations from multiple hosts, across a broad geographic range. Sequence characterised amplified region (SCAR) data identified an exotic host-generalist lineage and a native host-specialist lineage of O. minor, suggesting genetic structure in this species is defined by both host specificity and geography. In addition, host-range overlap, discordant tree topologies, and cryptic morphology indicate the presence of gene flow between alien races and endemic populations. Therefore, repeated introductions of alien races of O. minor from disparate sources leading to introgression with native populations, and cryptic race formation, seem to have contributed to the taxonomic confusion associated with this species. We speculate that radiations associated with broad host range and divergent host ecologies may have promoted the unusually wide geographic distribution and diversification of this species. Finally, evidence of multiple shifts to exotic hosts, coupled with the predicted northward shift in climatic suitability, identify the potential for range expansion in alien races of O. minor, which may threaten nationally scarce native taxa with genetic assimilation. Our phylogenetic analysis provides a framework for identifying host races in Orobanche with a view to setting conservation priorities.     

A putative role for fusaric acid in biocontrol of the parasitic angiosperm Orobanche ramosa

Fusarium spp. are ubiquitous fungi found in soil worldwide as both pathogenic and nonpathogenic strains. The signals leading to disease or the absence of disease are poorly understood. We recently showed that fusaric acid (FA), a nonspecific toxin produced by most Fusarium spp., could elicit various plant defense responses at 100 nM without toxic effect. In this study, we checked for the effect of FA on root and root hairs, probable first site of contact between the fungi and the host. Large FA concentrations reduce root and root-hair growth and induce a rapid transient membrane hyperpolarization, followed by a large depolarization, due to the inhibition of H(+)-ATPase currents. Nanomolar concentrations of FA induced only an early transient membrane hyperpolarization of root hairs compatible with the induction of a signal transduction pathway. FA at 10(-7) M failed to induce salicylic acid- and jasmonic acid/ethylene-dependent defense-related genes but inhibited the germination of the angiosperm parasite Orobanche ramosa in contact of FA-pretreated Arabidopsis thaliana seedlings. These data suggest that FA at nontoxic concentrations could activate signal transduction components necessary for plant-defense responses that could contribute to biocontrol activity of Fusarium spp.     

Cloning of a cryptochrome homologue from the holoparasitic plant Orobanche minor Sm.

Orobanche minor is a non-photosynthetic root holoparasitic plant. Although it is known that photosynthesis-related genes are inactivated or have been eliminated from the plastid genomes of holoparasites, little is known about the alterations in their genes involved in the signaling networks by which light regulates photosynthesis. Cryptochromes (crys), which are blue-light receptors, appear to control both photosynthesis-related and non-photosynthetic responses to light in higher plants. Because we are interested in to what extent a cry-mediated light signaling network remains in the holoparasites, we cloned CRY homologous cDNA from O. minor (OmCRY1) and used real-time RT-PCR to compare its expression under natural daylight and darkness. We found that the OmCRY1 has a high degree of homology with CRY1 s from photosynthetic plants. Expression of the OmCRY1 gene was higher in plants grown in the dark than that in the plants grown under natural daylight. This is the first report of the gene expression of a blue-light receptor in non-photosynthetic plants.     

Elevated atmospheric CO2 influences the interaction between the parasitic angiosperm Orobanche minor and its host Trifolium repens

The influence of the root holoparasitic angiosperm Orobanche minor Sm. on the biomass, photosynthesis, carbohydrate and nitrogen content of Trifolium repens L. was determined for plants grown at two CO₂ concentrations (350 and 550 μmol mol⁻¹). Infected plants accumulated less biomass than their uninfected counterparts, although early in the association there was a transient stimulation of growth. Infection also influenced biomass allocation both between tissues (infected plants had lower root:shoot ratios) and within tissues:infected roots were considerably thicker before the point of parasite attachment and thinner below. Higher concentrations of starch were also found in roots above the point of attachment, particularly for plants grown in elevated CO₂. Elevated CO₂ stimulated the growth of T. repens only during the early stages of development. There was a significant interaction between infection and CO₂ on growth, with infected plants showing a greater response, such that elevated CO₂ partly alleviated the effects of the parasite on host growth. Elevated CO₂ did not affect total O. minor biomass per host, the number of individual parasites supported by each host, or their time of attachment to the host root system. Photosynthesis was stimulated by elevated CO₂ but was unaffected by O. minor . There was no evidence of down-regulation of photosynthesis in T. repens grown at elevated CO₂ in either infected or uninfected plants. The data are discussed with regard to the influence of elevated CO₂ on other parasitic angiosperm-host associations and factors which control plant responses to elevated CO₂.     

Can source–sink relations explain responses of tobacco to infection by the root holoparasitic angiosperm Orobanche cernua?

The achlorophyllous holoparasitic angiosperm Orobanche cernua reduced biomass accumulation of its tobacco host, such that 73-d-old plants achieved 29% of the biomass of control plants. The difference in biomass between infected and uninfected tobacco could be accounted for directly by diversion of dry matter to the parasite. Thus, the smaller infected plants were responsible for the production of as much dry matter in host and parasite as their larger uninfected counterparts. The productivity of the infected system was maintained by: (a) sustained production of leaf area (a greater leaf area ratio); (b) increased specific leaf area, and (c) delayed senescence. When tobacco was inoculated with different densities of the parasite, the amount of dry matter accumulated by the parasite was not changed, suggesting that a finite amount of resource was available to the parasite. The response of the host to infection can be explained by simple source–sink interactions and the data are discussed with respect to other parasitic angiosperm–host systems which show different types of responses.     

Host‐driven divergence in the parasitic plant Orobanche minor Sm. (Orobanchaceae)

Many parasitic angiosperms have a broad host range and are therefore considered to be host generalists. Orobanche minor is a nonphotosynthetic root parasite that attacks a range of hosts from taxonomically disparate families. In the present study, we show that O. minor sensu lato may comprise distinct, genetically divergent races isolated by the different ecologies of their hosts. Using a three‐pronged approach, we tested the hypothesis that intraspecific taxa O. minor var. minor and O. minor ssp. maritima parasitizing either clover (Trifolium pratense) or sea carrot (Daucus carota ssp. gummifer), respectively, are in allopatric isolation. Morphometric analysis revealed evidence of divergence but this was insufficient to define discrete, host‐specific taxa. Intersimple sequence repeat (ISSR) marker‐based data provided stronger evidence of divergence, suggesting that populations were isolated from gene flow. Phylogenetic analysis, using sequence‐characterized amplified region (SCAR) markers derived from ISSR loci, provided strong evidence for divergence by clearly differentiating sea carrot‐specific clades and mixed‐host clades. Low levels of intrapopulation SCAR marker sequence variation and floral morphology suggest that populations on different hosts are probably selfing and inbreeding. Morphologically cryptic Orobanche taxa may therefore be isolated from gene flow by host ecology. Together, these data suggest that host specificity may be an important driver of allopatric speciation in parasitic plants.     

First record of potential bird pollination in the holoparasitic genus Orobanche L.

Pollinators play an important role in the reproduction of zoophilous plants. A shift in pollinators has often been observed for oceanic island plants, probably because of the differences in fauna. In this study, we obtained data on pollinator shifts from insects to birds in Orobanche boninsimae (Orobanchaceae), a holoparasitic plant species endemic to the Bonin (Ogasawara) Islands, oceanic islands in the Pacific. We observed pollination and measured seed viability in O. boninsimae and its continental sister species O. coerulescens. We found that two passerine birds, the Japanese white-eye (Zosterops japonicus) and bulbul (Hypsipetes amaurotis squameiceps), visited the flowers and sucked the nectar of O. boninsimae, while only insects visited those of O. coerulescens. Viable seeds were produced under pollinator-excluded treatments in the two Orobanche species, indicating that the seeds were produced by automatic self-pollination and/or apomixis. These results suggest that O. boninsimae may be pollinated by birds and can produce seeds by automatic self-pollination/apomixis. This is the first record of visitation of the genus Orobanche by birds. Studies of pollination systems in native plants on the Bonin Islands are few compared to those on other oceanic islands, and O. boninsimae may provide a valuable example of pollinator shifts in the Bonin Islands.     

寄生被子植物吸器的研究

从吸器的发生、结构、功能、寄生的生理机制等方面概述了寄生被子植物吸器的研究进展。     

Solute fluxes from tobacco to the parasitic angiosperm Orobanche cernua and the influence of infection on host carbon and nitrogen relations

Orobanche species are holoparasites which are very efficient sinks for host-derived solutes. Here, we report the use of direct measurements of xylem sap solute concentrations and water fluxes, together with a modelling procedure to calculate element fluxes within an association between Orobanche cernua and its tobacco host. Infection of tobacco by the parasite markedly influenced carbon acquisition and partitioning; net fixation of carbon was 20% higher in infected tobacco compared with controls. Orobanche cernua caused a 84% increase in net carbon flux moving downward from the tobacco shoot and 73% of this carbon was intercepted by the parasite, almost entirely through the phloem (>99%). Further, the parasite also exerted a large impact on the nitrogen relations of the plant, notably nitrate uptake was stimulated and the amino acid content of xylem sap was lower. The parasite also relied heavily on host phloem for the supply of other resources, with only 5 to 15% of N, and 16% of K, 23% of Na, 63% of Mg and 13% of S being derived from the xylem. Thus, we provide quantitative information on the phloem dependency of the parasite and show that host carbon and nitrogen metabolism is stimulated as a consequence of infection.     

Seedling diversity in Hydatellaceae: implications for the evolution of angiosperm cotyledons

BACKGROUND AND AIMS: Cotyledon number has long been a primary morphological feature distinguishing monocots from other angiosperms. Recent placement of Hydatellaceae near the early-divergent angiosperm order Nymphaeales, rather than in the monocot order Poales, has prompted reassessment of seedling morphology in this poorly known family. METHODS: Seedlings of six species representing all eco-geographical groups of Hydatellaceae are described using light and scanning electron microscopy. KEY RESULTS: Two seedling types were discovered. Material examined of Trithuria submersa, T. bibracteata, T. austinensis and T. filamentosa possess a transparent bilobed sheathing structure that surrounds the main axis below the first foliage leaf. The seed coat is attached to the sheathing structure. Seedlings of Trithuria lanterna and T. konkanensis lack a sheathing structure, and the seed coat is attached to a short, narrow lateral outgrowth on the main axis of the seedling. CONCLUSIONS: The sheathing structure that is present in seedlings of some Hydatellaceae could be homologized with the two united cotyledons of water lilies. It also resembles the single cotyledon of some monocots, and hence demonstrates a possible pathway of the origin of a monocot-like embryo, though no homology is implied. The sheathing structure is reduced in Trithuria lanterna and T. konkanensis, and the short, narrow outgrowth of its seedling could represent a single cotyledon. This synapomorphy suggests that the only Indian species of Hydatellaceae, T. konkanensis, is closer to the northern Australian T. lanterna than to the south-western Australian T. bibracteata.     

Localization of photosynthetic metabolism in the parasitic angiosperm Cuscuta reflexa

Cells capable of photosynthesis in the parasitic angiosperm Cuscuta reflexa Roxb. (dodder) are highly localized. Immunolocalization of ribulose-1,5 bisphosphate carboxylase-oxygenase (Rubisco) and autofluorescence of chlorophyll in transverse sections of stems showed that they were largely restricted to a band of cells adjacent to the vascular bundles, consequently, the concentrations of Rubisco and chlorophyll were low per unit area or fresh weight. When ¹⁴CO₂ was supplied to stem segments of C. reflexa it preferentially accumulated in these cells adjacent to the vasculature. Although the conductance for CO₂ movement to the cells containing chlorophyll and Rubisco was very low, both the light reactions and dark reactions of photosynthesis appeared to be functional. De-epoxidation of the xanthophyll-cycle pigments after exposure to high light, and the chlorophyll fluorescence parameters, photochemical quenching (qP), non-photochemical quenching (NPQ) and the quantum efficiency of photosystem II (φPSII) responded normally to changes in photon flux density, indicating functional light-driven electron transport. The response of CO₂ exchange to photon flux density followed a typical hyperbolic curve, and positive rates of CO₂ fixation occurred when external CO₂ was increased to 5%. We propose that CO₂ for carbon assimilation is derived from internally respired CO₂ and that this layer of photosynthetic cells makes a positive contribution to the carbon budget of C. reflexa. Received: 23 October 1997 / Accepted: 16 December 1997     

Confirmation and quantification of strigolactones, germination stimulants for root parasitic plants Striga and Orobanche, produced by cotton

The germination stimulants for root parasitic plants Striga and Orobanche produced by cotton (Gossypium hirsutum L.) were examined in detail. Seeds of cotton were germinated and grown on glass wool wetted with sterile distilled water in sterile filter units. The root exudate was collected daily and extracted with ethyl acetate. Each of these ethyl acetate extracts was analyzed directly by high-performance liquid chromatography linked with tandem mass spectrometry (LC/MS/MS). The results demonstrate that cotton roots exuded strigol and strigyl acetate, but no other known strigolactones such as orobanchol and alectrol. The production of strigol was detected even in the root exudate collected during the first 24 h of incubation and reached a maximum 5-7 days later. The average exudation of strigol and strigyl acetate during the incubation period was ca. 15 and 2 pg/plant/day, respectively, indicating that strigol mainly contributed to germination stimulation by the cotton root exudate.     

药用寄生植物菟丝子属,列当属和无根藤属氨基酸的分析

本文测定了菟丝子属、列当属和无根藤属某些种的种子和植株氨基酸的种类组成和含量。结果表明,3个属种子和植株氨基酸均在15种以上,且含量丰富,特别是必需氨基酸的含量较高。文中讨论了氨基酸的药用和在种子鉴定与化学分类上的作用,探讨了开发应用的前景。     

Specific developmental pathways underlie host specificity in the parasitic plant Orobanche

Parasitic angiosperms are an ecologically and economically important group of plants. However our understanding of the basis for host specificity in these plants is embryonic. Recently we investigated host specificity in the parasitic angiosperm Orobanche minor, and demonstrated that this host generalist parasite comprises genetically defined races that are physiologically adapted to specific hosts. Populations occurring naturally on red clover (Trifolium pratense) and sea carrot (Daucus carota subsp. gummifer) respectively, showed distinct patterns of host specificity at various developmental stages, and a higher fitness on their natural hosts, suggesting these races are locally adapted. Here we discuss the implications of our findings from a broader perspective. We suggest that differences in signal responsiveness and perception by the parasite, as well as qualitative differences in signal production by the host, may elicit host specificity in this parasitic plant. Together with our earlier demonstration that these O. minor races are genetically distinct based on molecular markers, our recent data provide a snapshot of speciation in action, driven by host specificity. Indeed, host specificity may be an underestimated catalyst for speciation in parasitic plants generally. We propose that identifying host specific races using physiological techniques will complement conventional molecular marker-based approaches to provide a framework for delineating evolutionary relationships among cryptic host-specific parasitic plants.Key words: host specificity, parasitic plant, broomrape, orobanche, speciation     

Correlated evolution of life history and host range in the nonphotosynthetic parasitic flowering plants Orobanche and Phelipanche (Orobanchaceae)

Most members of the nonphotosynthetic parasitic genera Orobanche and Phelipanche (Orobanchaceae) have narrow host ranges, and, as they grow on perennial hosts, are (at least potentially) perennial themselves. A few species, however, have wide host ranges and grow on annual hosts, and are thus (at least facultatively) annuals themselves. Among the latter are the weedy species, which include economically important pest taxa such as Orobanche crenata or Phelipanche aegyptiaca. Using a phylogenetically based maximum likelihood approach, which takes phylogenetic and branch length uncertainty into account, we can show that the life trait host range and life history evolve in a correlated fashion. This supports the hypothesis that parasite specialization is associated with predictable resources (i.e. long-lived hosts) and generalism with unpredictable ones (i.e. short-lived hosts), a pattern often found in animal parasites. The mechanisms and temporal sequence of the life trait changes and their interrelations remain speculative.