Eight polymorphic microsatellite markers for Rhinanthus minor

Little research has been carried out on the genetics of Rhinanthus minor to date. To enable study of this species, eight polymorphic microsatellite loci were developed, using a genomic library enriched for microsatellites. All loci are polymorphic in the two UK populations tested, Bardister and Oxwich Bay. These microsatellite markers will be useful for studying genetic structure and subspecies differences of R. minor.     

Differential resistance among host and non-host species underlies the variable success of the hemi-parasitic plant Rhinanthus minor

BACKGROUND AND AIMS: Rhinanthus minor is a root hemiparasitic plant that attacks a wide range of host species which are severely damaged by the parasite. Rhinanthus minor also attempts unsuccessfully to form connections to a range of non-hosts which in contrast are not damaged by the parasite; however, the underlying physiological basis of these differences is not fully understood. METHODS: Biomass of host-parasite combinations was studied, and histology, electron microscopy and FT-IR microspectroscopy were used to determine the cellular-level interactions between Rhinanthus haustoria (the parasite's connective structure) and the roots of a range of potential host species. RESULTS: Two distinct defence responses were observed in the non-host forbs Plantago lanceolata and Leucanthemum vulgare. Firstly, L. vulgare was able to encapsulate the parasite's invading structures preventing it from gaining access to the stele. This was supported by FT-IR microspectroscopy, used to monitor lignification in response to Rhinanthus haustoria. Secondly, host cell fragmentation was observed at the interface between the parasite and P. lanceolata. Growth data confirmed the non-host status of the two forbs whilst, in contrast, grasses and a legume which were good hosts showed no evidence of defence at the host/parasite interface. CONCLUSIONS: Variable resistance to Rhinanthus is shown for the first time to be controlled by cellular-level resistance to haustoria by either cell fragmentation or lignification at the host/parasite interface.     

Progress in parasitic plant biology: host selection and nutrient transfer

Host range varies widely among species of parasitic plants. Parasitic plants realize host selection through induction by chemical molecular signals, including germination stimulants and haustoria-inducing factors (HIFs). Research on parasitic plant biology has provided information on germination, haustorium induction, invasion, and haustorial structures and functions. To date, some molecular mechanisms have been suggested to explain how germination stimulants work, involving a chemical change caused by addition of a nucleophilic protein receptor, and direct or indirect stimulation of ethylene generation. Haustorium initiation is induced by HIFs that are generated by HIF-releasing enzymes from the parasite or triggered by redox cycling between electrochemical states of the inducers. Haustorium attachment is non-specific, however, the attachment to a host is facilitated by mucilaginous substances produced by haustorial hairs. Following the attachment, the intrusive cells of parasites penetrate host cells or push their way through the host epidermis and cortex between host cells, and some types of cell wall-degrading enzymes may assist in the penetration process. After the establishment of host-parasite associations, parasitic plants develop special morphological structures (haustoria) and physiological characteristics, such as high transpiration rates, high leaf conductance, and low water potentials in hemiparasites, for nutrient transfer and resource acquisition from their hosts. Therefore, they negatively affect the growth and development and even cause death of their hosts.     

Interactions between arbuscular mycorrhizal fungi and the hemiparasitic angiosperm Rhinanthus minor during co-infection of a host

The outcome of dual infection of the grass Lolium perenne L. by arbuscular mycorrhizal (AM) fungi and the parasitic angiosperm Rhinanthus minor L. was investigated in a glasshouse study. Colonization of L. perenne roots by AM fungi was significantly reduced by the presence of R. minor , as was host growth which fell by 44–51%. It was concluded that these two responses were linked, with AM colonization declining in response to the reduction in availability of host carbon. Parasite growth and reproductive output rose by 58% and 47% respectively when the hosts were mycorrhizal. These trends were unrelated to the attachment success of the parasite, but were accompanied by a significant increase in the formation of secondary haustoria. The benefits afforded the parasite when the hosts were mycorrhizal were attributed to increased carbon and nutrient flux resulting from alternations in sink strength. Host responses to parasitism and mycorrhizal colonization were not affected by the interaction between the two symbionts. However, the suggestion is made that the interaction between the AM fungi and parasite might have long-term ecological implications for the host species via its impact on parasite fecundity.     

Growth, gas exchange and water use efficiency of the facultative hemiparasite Rhinanthus minor associated with hosts differing in foliar nitrogen concentration

The facultative hemiparasitic angiosperm Rhinanthus minor was grown on 11 different host species, and in the absence of a host. The height of R. minor parasitising the legume Trifolium pratense exceeded that of unattached plants by more than an order of magnitude, with performance on grasses and non-legume dicotlyedonous hosts between these two extremes. Light saturated rates of photosynthesis in R. minor on different hosts ranged from 1. 5 to 22. 5 μmol g⁻¹ dry weight min⁻¹, and were positively correlated with growth. Foliar nitrogen concentrations in the parasite exceeded those of the hosts. The former were positively related to light saturated rates of photosynthesis, and inversely related to photosynthetic nitrogen use efficiency. There was no relationship between leaf nitrogen concentration and water use efficiency in R. minor. The data are discussed in relation to studies of nitrogen and water use in mistletoes.     

The host range and selectivity of a parasitic plant: Rhinanthus minor L.

Summary Rhinanthus minor (Yellow-rattle) is a widespread hemiparasitic plant of grassland habitats throughout Britain. Association analysis of the dune vegetation at Holme-next-the-Sea in eastern England revealed only two potential host plants through positive association. In contrast direct examination of the root systems revealed haustorial connections with 20 host species. The number of species parasitized by one plant ranged from one to seven. Data from another four sites in Britain and one in central Europe indicate that the natural host range of R. minor encompasses at least 50 species from 18 families with 22% in the Leguminosae and 30% in the Gramineae. Comparison of the number of haustorial connections made to each species with the abundance of roots in the soil shows that R. minor is a highly selective parasite, but that the selectivity is not consistent between populations or between plants from different parts of the same population. The reasons for host selectivity are discussed.     

Genome and plastome effects on photosynthesis parameters in Oenothera species of differential low-temperature tolerance

The influence of the plastid genome (plastome) on the temperature dependence of chlorophyll fluorescence parameters was studied in four Oenothera species from climatically different habitats. Equipped with their natural plastome, the species could be arranged with respect to their low-temperature tolerance in the order Oe'villosa' (plastome I) = Oe argillicola (V) > Oe. grandiflora (III) > Oe elata ssp. hookeri (J). Exchanging the naturally occurring plastome with those from other species did not result in changes of the photosynthetic performance of the genotypes, except for decreasing pigment contents and photosynthesis rates (on area basis) in Oe. elata ssp. hookeri with plastome III. Furthermore, plastome exchange in Oe. elata ssp. hookeri and in Oe. grandiflora did not affect kinetic properties of purified ribulose-l,5-bis-phosphate carboxylase/oxygenase (EC 4,1.1.39), indicating that the gene for the large subunit of this enzyme has remained conserved during plastome evolution in Oenothera . It is concluded that the evolution of the plastome in North American Oenothera spp. did not influence the temperature adaptation of the photosynthetic apparatus, but that the latter is governed by effects residing in the nuclear genome.     

Impacts of Infection by Parasitic Angiosperms on Host Photosynthesis

Abstract: Parasitic angiosperms are a taxonomically diverse group of plants that invade host tissues and remove resources via a specialized structure known as the haustorium. Through the haustorium, carbon, water and mineral nutrients are withdrawn, often at the expense of host growth and vigour. In addition to the removal of resources from host plants, many parasitic angiosperms are also able to impact host growth via effects on host photosynthesis. In this paper we review what is known about how parasitic angiosperms affect host photosynthesis and the impact this has on host productivity. Holoparasites, that lack chlorophyll, act as extra sinks for host photosynthates and generally either enhance or have a neutral effect on host photosynthetic productivity. In contrast, hemiparasites, that are capable of some autotrophic carbon fixation, usually have a negative impact on host photosynthesis. Irrespective of the outcome of infection, the mechanisms involved in altering host photosynthesis are diverse and may act at either the leaf or whole-plant level. In some cases, parasites impact directly on host photosynthetic metabolism, while in others the effects are more indirect, for example through changing host architecture.     

A method for measuring whole plant photosynthesis in Arabidopsis thaliana

Measurement of photosynthesis of intact leaves of Arabidopsis thaliana has been prohibitive due to the small leaf size and prostrate growth habit. Because of the widespread use of Arabidopsis for plant science research it is important to have a procedure for accurate, nondestructive measurement of its photosynthesis. We developed and tested a method for analysis of photosynthesis in whole plants of Arabidopsis. Net carbon assimilation and stomatal conductance were measured with an open gas exchange system and photosynthetic oxygen evolution was determined from chlorophyll fluorescence parameters. Individual plants were grown in 50 cubic centimeter tubes that were attached with an air tight seal to an enclosed gas exchange chamber for measurement of carbon dioxide and water exchange by the whole plant. Chlorophyll fluorescence from intact leaves was simultaneously measured with a pulse modulated fluorometer. Photosynthetic CO₂ assimilation and stomatal conductance rates were calculated with established gas exchange procedures and O₂ evolution was determined from chlorophyll fluorescence measurement of Photosystem II yield. Carbon assimilation and oxygen evolution in response to light intensity and ambient CO₂ concentration was measured and is presented here to demonstrate the potential use of this method for investigation of photosynthesis of Arabidopsis plants in controlled environment conditions.     

寄生植物种子萌发特异性及其与寄主的识别机制

寄生植物广泛分布于不同的生态环境中,并具有不同的生育习性及与寄主识别特性.本文阐述了根寄生植物列当属和独脚金属种子萌发的特异性,以及目前已发现的寄生植物种子萌发的信号物质,并就不同萌发信号物质、植物激素、真菌代谢物在寄生植物种子识别寄主中的作用以及寄生植物种子预培养阶段的呼吸作用特性与萌发信号物质的活化机理等做了综述.探讨了各种列当不同分化类型的愈伤组织诱导、离体无菌侵染新系统及其在寄生植物与寄主互作识别研究中的应用,提出了寄生植物与寄主识别机理研究中存在的问题并对研究前景进行了展望.     

The action of a range of supplementary ultraviolet (UV) wavelengths on photosynthesis in Brassica napus L. in the natural environment: effects on PS II,CO2 assimilation and level of chloroplast proteins

The effects of different wavebands of UV radiation on photosynthesis and the expression and abundance of photosynthetic proteins in oilseed rape (Brassica napus L. cv. Rebel) were investigated. Plants were grown outdoors under natural radiation (52° N, 0° E) supplemented with six wavebands of UV radiation (0.4 Wm⁻²) between 313 nm and 356 nm. A control treatment was centred at 343 nm. Exposure to supplementary UV-A radiation (320–400 nm) had no significant effects, however UV-B radiation, centred at 313 nm, caused a marked reduction in photosynthesis. This decrease was related to a reduction in the initial carboxylation velocity of Rubisco which was further correlated with a large reduction in the expression and abundance of both large and small subunits of Rubisco. These results indicate a molecular mechanism behind UV-B induced reductions in photosynthesis per unit area in plants grown under field conditions. This revised version was published online in June 2006 with corrections to the Cover Date.     

The impact of phosphorus on interactions of the hemiparasitic angiosperm <Emphasis Type="Italic">Rhinanthus minor</Emphasis> and its host <Emphasis Type="Italic">Lolium perenne</Emphasis>

The effects of phosphorus supply on the outcome of interactions between the hemiparasitic angiosperm Rhinanthus minor L. with its host species Lolium perenne L. were investigated in a glasshouse experiment. Host plants were grown in 3-l pots in the presence and absence of R. minor at limiting (0.13 mm P) and optimal (0.65 mm P) concentrations of phosphorus for the growth of the host species. Phosphorus was supplied at 2-day intervals in the form of half-strength Long Ashton nitrate-based solution with phosphorus concentrations adjusted accordingly. Parasitism by R. minor significantly suppressed host growth, with final biomass losses ranging between 32% and 44%. Phosphorus supply had a marked impact on the outcome of the host-parasite interaction. By the end of the growing period, parasite biomass at 0.65 mm P was 90% lower than that achieved at 0.13 mm P. In contrast, host biomass at 0.65 mm P was 74% higher than achieved at 0.13 mm P, indicting that the negative impact of parasitism on the host species was reduced when phosphorus supply was increased. The effects of phosphorus on the host-parasite association appeared to be mediated by changes in both the morphological characteristics of the host roots and the relative sink strengths of the host and parasite. Received: 29 May 1999 / Accepted: 20 December 1999     

脱落酸对低温下雷公藤幼苗光合作用及叶绿素荧光的影响

以1年生雷公藤扦插苗为试材,研究低温胁迫下不同浓度外源脱落酸（ABA,0、5、10、15、20、25 mg·L^-1)叶面喷施处理对雷公藤叶片光合作用及叶绿素荧光参数的影响.结果表明：喷施20 mg·L^-1的ABA能显著提高雷公藤幼苗的抗冷性,减缓低温下雷公藤叶片净光合速率(P_n)、蒸腾速率(T_r)、气孔导度(g_s)、胞间CO₂浓度(C_i)的下降幅度,提高幼苗叶片的光合能力.低温处理6 d后,随着ABA浓度上升,雷公藤叶片的初始荧光(F_o)下降,最大荧光(F_m)和PSII最大光化学效率(F_v/F_m)上升,PSII实际光化学量子产量(Φ_PSⅡ)、光化学猝灭系数(q_P)先下降后上升,而非光化学猝灭系数(q_N)呈下降－上升－下降趋势.P_n、g_s、q_P、F_m和F_v/F_m均在20 mg·L^-1ABA处理时达到峰值.不同浓度ABA的相对电子传递速率(rETR)随着光化光强度增加呈先上升后下降的趋势,当光化光强度(PAR)达到395 μmol·m^-2s^-1时,各处理的rETR达到最高值,其中25 mg·L^-1和20 mg·L^-1ABA处理分别比对照高17.1%和5.2%.雷公藤叶片Φ_PSⅡ的光响应曲线均随光化光强度升高而下降,q_N的光响应曲线则呈相反趋势.     

Effects of long-term exposure to elevated UV-B radiation on the photosynthetic performance of five broad-leaved tree species

As part of an ongoing investigation into the effects of long-term UV-B radiation exposure on the growth and morphology of woody perennials, the gas exchange and photosynthesis of five common deciduous tree species were measured. All five tree species had been exposed to UV-B radiation for 5 years, in the field, at an enhancement level equivalent to an 18% ozone depletion. Measurements made during the fifth year of UV-B irradiation recorded reductions in light-saturated photosynthesis, transpiration and water use efficiencies. These changes were accompanied by marked reductions in individual leaf areas, stomatal density, stomatal conductance and carboxylation efficiency. There were no significant changes in the maximum variable fluorescence ratio, the quantum requirement of oxygen evolution, or light-saturated O₂ production. Analysis of the response of net carbon assimilation to changing intercellular CO₂ concentration (A/c_i response) demonstrated no significant change in stomatal limitation. Reductions in photosynthesis were consistent with decreased carboxylation efficiency. Although all five tree species were similarly affected by UV-B radiation treatment, the magnitude of the responses was species-specific. These findings demonstrate the need for more long-term experimentation and also suggest that changes in water use efficiency may be a significant factor in plants' responses to UV-B radiation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Like herbivores,parasitic plants are limited by host nitrogen content

Herbivores generally benefit from increased plant nitrogen content, because the nitrogen content of animals is much higher than that of plants. Consequently, high plant nitrogen alleviates the profound stoichiometric imbalance that herbivores face in their diets. Parasitic plants provide the opportunity to test this generalization for consumers across kingdoms. We fertilized two microhabitats in a California salt marsh that were dominated by Salicornia virginica or a mixture of S. virginica and Jaumea carnosa. The nitrogen content of both host plants and of the holoparasite Cuscuta salina (dodder) increased in fertilized plots in both microhabitats. Cuscuta preferred to attack Jaumea, although Jaumea had lower nitrogen content than Salicornia. When host nitrogen content was altered by fertilizing plots, however, the percent cover of the parasite doubled. Although parasitic plants and their hosts have similar tissue nitrogen contents, suggesting no stoichiometric imbalance between host and consumer, parasitic plants do not feed on host tissue, but on host xylem and phloem, which are very low in nitrogen. Consequently, parasitic plants face the same dietary stoichiometric constraints as do herbivores, and both herbivores and holoparasitic plants may respond positively to increases in host nitrogen status.     

Parasite–grass–forb interactions and rock–paper– scissor dynamics: predicting the effects of the parasitic plant Rhinanthus minor on host plant communities

1.  Parasitic plants affect the growth, reproduction and metabolism of their hosts and may also influence the outcome of competitive interactions between host species and, consequently, the structure of entire host communities.
2.  We investigate the effect of the root hemiparasitic plant Rhinanthus minor on plant community dynamics using a spatial theoretical model. The model is parameterized with data from pairwise interaction experiments under two nutrient levels between the hemiparasite and three grass species ( Cynosurus cristatus , Festuca rubra and Phleum bertolonii ) and three forb species ( Leucanthemum vulgare , Plantago lanceolata and Ranunculus acris ).
3.  Relative interaction coefficients were intransitive, with the dynamics of the system conforming to a rock–paper–scissors game. Stable deterministic dynamics emerge from parameters obtained under low-nutrient conditions. Under high-nutrient conditions, the dynamics are unstable, but are stabilized in spatially explicit models. The outcomes are sensitive to initial spatial pattern and frequency.
4.   Synthesis . This study supports the idea that hemiparasite populations may form 'shifting clouds' in natural populations and explains seemingly unpredictable shifts in host community structure following introduction of hemiparasites. Management of plant communities using hemiparasites needs to take these complex dynamics into account.