Morphological and molecular confirmation of Albugo resedae (Albuginales; Oomycota) as a distinct species from A. candida

The genus Albugo s.str. causes white blister rust on four families of the Brassicales, Brassicaceae, Capparaceae, Cleomaceae, and Resedaceae. Recent phylogenetic studies have revealed that several host specific lineages are present within Albugo on Brassicales, while it was also confirmed that Albugo candida has an exceptionally wide host range which extends from Brassicaceae to Cleomaceae and Capparaceae. The Albugo species infecting the Resedaceae was attributed in monographic studies as well as local floras to either A. resedae or, applying a broader species concept, to A. candida. In the present study, A. resedae specimens were morphologically and molecularly compared to the five Albugo species so far confirmed from Brassicales, A. candida, A. koreana, A. laibachii, A. lepidii, and A. voglmayrii. Both morphological differences of oospore ornamentation and phylogenetic analysis of cox2 mtDNA sequences provided evidence that A. resedae is distinct from A. candida and from the additional four species so far described from Brassicaceae. It thus seems possible that so far unknown factors restrict Albugo candida to Brassicaceae and its sister families, Cleomaceae and Capparaceae.     

A new presumably widespread species of Albugo parasitic to Strigosella spp. (Brassicaceae)

White blister rust is one of the most common diseases in Brassicaceae. Recently, molecular approaches revealed that apart from Albugo candida, several other more specialized species of the genus are causing this disease on Brassicaceae and the diversity of this group still remains largely unexplored. All newly described species have so far been sampled only from a limited geographic range, except for Albugo species which followed their invasive host from Europe to other continents. In this study we show that a previously unknown species of Albugo is causing white blister rust disease on two species of Strigosella. This species can be distinguished from other species of the genus Albugo both by its phylogenetic position and its unique oospore ornamentation that might be an adaptation to the harsh environment of the host plants. As a consequence, Albugo arenosa is described and illustrated as a new species, so far known from Strigosella africana in Iran and Spain, and Strigosella brevipes in Iran. Apart from Albugo candida and Albugo lepidii, this is the third hitherto known species of Albugo s. str. with a confirmed native distribution range of several thousand kilometres.     

Phylogeny of an Albugo sp. infecting Barbarea vulgaris in Denmark and its frequency of symptom development in natural populations of two evolutionary divergent plant types

The oomycete Albugo candida has long been considered a broad spectrum generalist pathogen, but recent studies suggest that it is diverged into several more specialized species in addition to the generalist Albugo candida sensu stricto. Whereas these species cause the disease white blister rust in many crucifer plants, asymptomatic endophytic infections may be important in the epidemiology of others. One of the plant species attacked by Albugo sp. is the wild crucifer Barbarea vulgaris ssp. arcuata, which is diverged into two phytochemically and genetically different types with different geographical distributions in Europe. These were previously shown to differ strongly in propensity to develop white rust upon controlled infections in the greenhouse. Here, we analyse the phylogenetic relatedness of this local Albugo sp. field isolate to other species and lines of Albugo spp., including others collected on B. vulgaris. We further ask whether the difference in incidence of white rust between the two types of B. vulgaris are also expressed in natural populations.     

中国白锈菌科分类研究 Ⅱ.爵床科上一新种及十字花科上的已知种

本文报道爵床科(Acanthaceae)上的尖药草白锈新种(Albugo aechmantherae Zhang etY.X.Wang sp.nov.)和十字花科(Cruciferae)上已知种白锈菌[A.candida(Pers.)O.Kuntze]及大孢白锈[A.macrospora(Togashi)S.Ito]的寄主新记录和寄主植物国内新记录。新种有汉文和拉丁文描述及形态图。     

Basic compatibility of Albugo candida in Arabidopsis thaliana and Brassica juncea causes broad-spectrum suppression of innate immunity

A biotrophic parasite often depends on an intrinsic ability to suppress host defenses in a manner that will enable it to infect and successfully colonize a susceptible host. If the suppressed defenses otherwise would have been effective against alternative pathogens, it follows that primary infection by the "suppressive" biotroph potentially could enhance susceptibility of the host to secondary infection by avirulent pathogens. This phenomenon previously has been attributed to true fungi such as rust (basidiomycete) and powdery mildew (ascomycete) pathogens. In our study, we observed broad-spectrum suppression of host defense by the oomycete Albugo candida (white blister rust) in the wild crucifer Arabidopsis thaliana and a domesticated relative, Brassica juncea. A. candida subsp. arabidopsis suppressed the "runaway cell death" phenotype of the lesion mimic mutant lsd1 in Arabidopsis thaliana in a sustained manner even after subsequent inoculation with avirulent Hyaloperonospora arabidopsis (Arabidopsis thaliana downy mildew). In sequential inoculation experiments, we show that preinfection by virulent Albugo candida can suppress disease resistance in cotyledons to several downy mildew pathogens, including contrasting examples of genotype resistance to H. arabidopsis in Arabidopsis thaliana that differ in the R protein and modes of defense signaling used to confer the resistance; genotype specific resistance in B. juncea to H. parasitica (Brassica downy mildew; isolates derived from B. juncea); species level (nonhost) resistance in both crucifers to Bremia lactucae (lettuce downy mildew) and an isolate of the H. parasitica race derived from Brassica oleracea; and nonhost resistance in B. juncea to H. arabidopsis. Broad-spectrum powdery mildew resistance conferred by RPW8 also was suppressed in Arabidopsis thaliana to two morphotypes of Erysiphe spp. following pre-infection with A. candida subsp. arabidopsis.     

白锈菌一新种

本文报道了在四川采集的寄生于荨麻科植物粗齿冷水花(Pilea fasciata Franch)叶片上白锈菌的一个新种,命名为冷水花白锈(Albugo pileae rao et Y. Qin sp. nov.),并附拉丁文和汉文描述及形态图,还讨论了新种与近似种之间的区别。     

A new perspective on the evolution of white blister rusts: <Emphasis Type="Italic">Albugo</Emphasis> s.str. (Albuginales; Oomycota) is not restricted to Brassicales but also present on Fabales

For almost all groups of pathogens, unusual and rare host species have been reported. Often, such associations are based on single or few collections only, which are frequently hard to access. Many of them later prove to be due to misidentification of the host, the pathogen, or both. Therefore, such reports are often disregarded, or treated anecdotally in taxonomic and phylogenetic studies, regardless of their potential importance to unravelling the evolution of the entire group. Concerning oomycete biotrophs there are several reports of unusual and rare hosts for hardly known pathogens. In the order Fabales, for example, a single species of Albugo, A. mauginii, was described as parasitic to Onobrychis crista-galli about 80 years ago, but not recorded again. All other confirmed members of Albugo s.str. are parasitic to representatives of the families Brassicaceae, Capparaceae, Cleomaceae, and Resedaceae in the order Brassicales. In the present study, molecular phylogenetic analysis of cox2 mtDNA sequences and morphological investigations on an original specimen confirmed the occurrence of a member of Albugo on Fabaceae hosts, with the characteristic thin wall of the secondary sporangia, which is almost uniform in thickness. In phylogenetic analyses the species results as embedded within Albugo s.str. Therefore, it is concluded that the natural host range of Albugo s.str. extends from Brassicales to Fabales via host jumping. Our results underscore that unrevised reports of pathogens from unusual hosts should be reconsidered carefully to obtain a more complete picture of pathogen diversity and evolution.     

A comparison of Peronospora parasitica (Downy mildew) isolates from Arabidopsis thaliana and Brassica oleracea using amplified fragment length polymorphism and internal transcribed spacer 1 sequence analyses

Amplified fragment length polymorphism (AFLP) fingerprints and internal transcribed spacer 1 (ITS1) sequences from 27 Peronospora parasitica isolates (collected from Arabidopsis thaliana or Brassica oleracea), 5 Albugo candida isolates (from the same hosts and from Capsella bursa-pastoris), and 1 Bremia lactucae isolate (from Lactuca sativa) were compared. The AFLP analysis divided the isolates into five groups that correlated with taxonomic species and, in most cases, with host origin. The only exception was a group consisting of A. candida isolates from both B. oleracea and C. bursa-pastoris. ITS1 sequence analysis divided the isolates into the same five groups, demonstrated the divergence between P. parasitica isolates from A. thaliana and B. oleracea, and, using previously published ITS1 sequences, clearly showed the relationship between A. candida isolates from different hosts.     

辣根属和豆瓣菜属(十字花科)系统位置的分子证据

对十字花科葶苈族的辣根属、南芥族的豆瓣菜属及相关属种植物的叶绿体DNA的trnL内含子和trnL-F基因间隔区序列进行了测定分析。结果表明,辣根属植物与南芥族的山芥属、蔊菜属、豆瓣菜属、碎米荠属在系统发育树中聚成一支,与葶苈族的模式属葶苈属植物相隔较远,结合形态特征,本研究认为辣根属应从葶苈族移出,其系统位置应靠近山芥属、蔊菜属、豆瓣菜属、碎米荠属植物;此外,系统发育树中,豆瓣菜属植物并入碎米荠属中,表明二者具有更近的亲缘关系,本研究结果不支持《中国植物志》第33卷对辣根属和豆瓣菜属的系统位置的处理。     

Genetic diversity within the Albugo candida complex (Peronosporales, Oomycota) inferred from phylogenetic analysis of ITS rDNA and COX2 mtDNA sequences

Albugo candida is a destructive fungus infecting brassicaceous hosts. The genetic diversity within the A. candida complex from various host plants was investigated by sequence analysis of the internal transcribed spacer (ITS) region of rDNA and the cytochrome c oxidase subunit II (COX2) region of mtDNA. The aligned nucleotide sequences of A. candida shared significantly high distances, up to 20.4 and 8.9%, in two genes. The phylogenetic trees, obtained using the Bayesian method and maximum parsimony analysis, showed two separate groups that corresponded to the host genera. Group I included A. candida isolates infecting Arabis, Autrieta, Berteroa, Biscutella, Brassica, Cardaminopsis, Diplotaxis, Eruca, Erysimum, Heliophila, Iberis, Lunaria, Raphanus, Sinapis, Sisymbrium, and Thlaspi. Group II contained all isolates from Capsella, Descurainia, Diptychocarpus, Draba, and Lepidium. The genetic similarities between the two genes among isolates within Group I were 99.0-100% and 99.6-100%, while those within Group II were 90.4-100% and 91.1-100%, respectively, showing considerably lower values than for Group I. The A. candida isolates from Capsella bursa-pastoris in Korea are clearly separated by sequence analysis for the two genes compared to those from Wales, England, and the USA. Based on the molecular data from the two genes, we suggest the high degree of genetic diversity exhibited within A. candida complexes warrants their division into several distinct species.     

中国白锈菌科分类研究——Ⅲ.茄科上一新种及苋科上的已知种

本文报道茄科(Solanaceae)上的天仙子白锈新种(Albugo hyoscyami Zhang, Y.X.Wang et Fu sp.nov.)和苋科(Amaranthaceac)上已知种苋白锈[A.bliti(Biv.)O.Kuntze]及牛膝白锈[A.achyranthis(P.Henn.)Miyabe]的寄主新记录和地理新分布。新种有汉文和拉丁文描述及形态图。     

White butterfly (

Once widespread, Cook?s scurvy grass (or nau, Lepidium oleraceum) is now confined to a few offshore populations. Classed as nationally endangered by the New?Zealand Department of Conservation, populations of Cook?s scurvy grass are threatened by a number of factors, including introduced herbivorous insect species such as the white butterfly (Pieris rapae) and white rust infection caused by the oomycete Albugo candida. In this paper, we investigate the occurrence of white butterfly on Cook?s scurvy grass and possible interactions with the white rust infection on the northernmost of the Matariki Islands in the Firth of Thames, New?Zealand. We found that larger host plants were more likely to be infested with white butterfly. The occurrence of white butterfly eggs and larvae also decreased as levels of white rust increased. Twenty-eight percent of the white butterfly larvae collected and reared in the laboratory were parasitised by the braconid wasp species Cotesia rubecula. We also reared a hyperparasitoid belonging to the super-family Chalcidoidea from one of the parasitoid cocoons. Further studies on the trophic interactions between Cook?s scurvy grass, Albugo?candida and white butterfly and its parasitoids could improve the understanding of the threats posed by plant pathogens and insect herbivores to populations of Cook?s scurvy grass, which in turn may lead to new management strategies for conservation.     

The only known white blister rust on a basal angiosperm is a member of the genus <Emphasis Type="Italic">Albugo</Emphasis>

Rare pathogens on unusual hosts are often providing valuable insight into the evolution of the pathogen group concerned, but it is often challenging to obtain sequence data for these, as because only very few, often decades-old specimens are available. One such example is Albugo tropica, the white blister pathogen of a basal angiosperm in the genus Peperomia (Piperaceae). For this species, only two, more than 70 and over 120-year-old collections available. Here, sequence data for A. tropica are reported and phylogenetic reconstructions reveal it as the sister group to all other white blister rusts of the genus Albugo. Its isolated position is also reflected by several morphological differences to the other species of the genus, such as very thin-walled sporangia and almost smooth oospores. The isolated phylogenetic position of the pathogen and its host might indicate that it is a relict species trapped on its host. The sister-group relationship to all members of the genus Albugo s.str., which have been investigated using molecular phylogenetics, hints at the possibility, that Albugo might have originated in South America or Gondwana and has later radiated in the holarctic on members of the Brassicales.     

Specificity of a Y-linked gene in the flea beetle Phyllotreta nemorum for defences in Barbarea vulgaris

A Y-linked gene (R-gene) in the flea beetle Phyllotreta nemorum L. (Coleoptera: Chrysomelidae: Alticinae) confer the ability of larvae to survive on types of the plant Barbarea vulgaris R.Br. (Brassicaceae) which are immune to attack by susceptible conspecifics. Two near-isogenic flea beetle lines were developed. The YE-line contained the Y-linked R-gene, and male larvae from this line survived on B. vulgaris. The ST-line did not contain the gene and did not survive on the plant. The YE-line had been developed through 8–9 generations of backcrosses (YE-males with ST-females) and the two lines were considered to be isogenic except for genes located on the Y-chromosome.A single copy of the Y-linked gene is sufficient to transfer a susceptible genotype (ST) into a resistant genotype (YE) which is able to utilize a plant that is immune to attack by specimens without R-genes. The Y-linked gene had no effects on survival on other plant species tested. The gene did not have any effect on developmental times and weights of adult beetles reared on other plants than B. vulgaris. Developmental times of larvae with the Y-linked gene were longer on B. vulgaris than on normal host plants, R. sativus and S. arvensis, but the adults obtained the same size on these plant species. No trade-offs of the Y-linked gene were discovered. The results suggest that the occurrence of the Y-linked gene is a derived trait which has enabled the flea beetle to expand its host plant range. The evolution of a host shift to B. vulgaris seems not to be favoured by the presence of this single gene.     

Differing acceptance of familiar and unfamiliar plant species by an oligophagous beetle

The mustard leaf beetle, Phaedon cochleariae (F.) (Coleoptera: Chrysomelidae), is specialized to feed and develop on various species within the Brassicaceae. In this study, we investigated the acceptance of several host plant species (Brassica rapa L. and Sinapis alba L.), commonly used by the beetle (familiar plants), and of various unfamiliar plants, including systematically and chemically related [Bunias orientalis L. (Brassicaceae) and Tropaeolum majus L. (Tropaeolaceae), both Brassicales], as well as unrelated non‐host plant species [Plantago lanceolata L. (Plantaginaceae); Lamiales]. Emphasis was laid on the acceptance of the neophyte B. orientalis, and on underlying cues responsible for the acceptance of the various species. Behavioural responses to plant volatiles were studied using a static four‐chamber olfactometer. Stimulants and deterrents were investigated by bioassay‐guided solid phase extraction and semi‐preparative high performance liquid chromatography. A difference in acceptance of plant species was found: odours and polar compounds of all Brassicales evoked attraction and feeding stimulation, respectively, in Ph. cochleariae. Glucosinolates and their volatile hydrolysis products could be the main compounds that are involved in attraction of the beetles. In contrast, Ph. cochleariae did not respond to odours of the non‐host P. lanceolata, and some fractions of this plant had feeding‐deterrent effects, due to the presence of iridoid glycosides, among others. Although adult females accepted the neophyte B. orientalis for oviposition, neonate larvae did not survive on it. The flavonoid‐containing fraction of this plant was deterrent, whereas a similar fraction had been shown to cause some feeding stimulation when derived from S. alba. Differences in qualitative and quantitative composition of related metabolites lead to differentiated plant acceptance, proving the complexity of plant cues and of insect responses that determine host acceptance behaviour. The possibility of a diet breadth enlargement to B. orientalis and the role of Ph. cochleariae as a putative native biocontrol agent of this invasive plant are discussed.     

WRR4, a broad-spectrum TIR-NB-LRR gene from Arabidopsis thaliana that confers white rust resistance in transgenic oilseed brassica crops

White blister rust caused by Albugo candida (Pers.) Kuntze is a common and often devastating disease of oilseed and vegetable brassica crops worldwide. Physiological races of the parasite have been described, including races 2, 7 and 9 from Brassica juncea , B. rapa and B. oleracea , respectively, and race 4 from Capsella bursa-pastoris (the type host). A gene named WRR4 has been characterized recently from polygenic resistance in the wild brassica relative Arabidopsis thaliana (accession Columbia) that confers broad-spectrum white rust resistance ( WRR ) to all four of the above Al. candida races. This gene encodes a TIR-NB-LRR (Toll-like/interleukin-1 receptor-nucleotide binding-leucine-rich repeat) protein which, as with other known functional members in this subclass of intracellular receptor-like proteins, requires the expression of the lipase-like defence regulator, enhanced disease susceptibility 1 ( EDS1 ). Thus, we used RNA interference-mediated suppression of EDS1 in a white rust-resistant breeding line of B. napus (transformed with a construct designed from the A. thaliana EDS1 gene) to determine whether defence signalling via EDS1 is functionally intact in this oilseed brassica. The eds1-suppressed lines were fully susceptible following inoculation with either race 2 or 7 isolates of Al. candida. We then transformed white rust-susceptible cultivars of B. juncea (susceptible to race 2) and B. napus (susceptible to race 7) with the WRR4 gene from A. thaliana . The WRR4-transformed lines were resistant to the corresponding Al. candida race for each host species. The combined data indicate that WRR4 could potentially provide a novel source of white rust resistance in oilseed and vegetable brassica crops.     

Alyssum mughlaei (Brassicaceae), a new species from Southwest Anatolia

Alyssum mughlaei (Brassicaceae) is a new species, described and illustrated from Southwest Anatolia, Turkey.     

A new species of <Emphasis Type="Italic">Pustula</Emphasis> (Oomycetes,Albuginales) is the causal agent of sunflower white rust

White blister rust of sunflower is an emerging disease that is among the most important diseases in this crop in South Africa and has recently spread to Europe. For the genus Albugo, it has been demonstrated that species are mostly at least host genus specific and that several previously overlooked species are present on Brassicaceae. It thus seems likely that previously unrecognised species are also present in the genus Pustula. Based on previous phylogenetic reconstruction in combination with differences in oospore ornamentation, it is revealed that Pustula on sunflower, previously attributed to Pustula tragopogonis (syn. Albugo tragopogonis), is distinct from Pustula on Tragopogon. Therefore, this pathogen is described as a new species, Pustula helianthicola. In addition, taxonomic observations revealed that Pustula tragopogonis is an incorrect name and is replaced by the new combination, Pustula obtusata.