Molecular phylogeny of centrohelid heliozoa,a novel lineage of bikont eukaryotes that arose by ciliary loss

Abstract Recent molecular and cellular evidence indicates that eukaryotes comprise three major lineages: the probably ancestrally uniciliate protozoan phylum Amoebozoa; the ancestrally posteriorly uniciliate opisthokont clade (animals, Choanozoa, and fungi); and a very diverse ancestrally biciliate clade, the bikonts—plants, chromalveolates, and excavate and rhizarian Protozoa. As Heliozoa are the only eukaryote phylum not yet placed on molecular sequence trees, we have sequenced the 18S rRNA genes of three centrohelid heliozoa, Raphidiophrys ambigua, Heterophrys marina, and Chlamydaster sterni, to investigate their phylogenetic position. Phylogenetic analysis by distance and maximum likelihood methods allowing for intersite rate variation and invariable sites confirms that centrohelid heliozoa are a robust clade that does not fall within any other phyla. In particular, they are decisively very distant from the heterokont pedinellid chromists, at one time thought to be related to heliozoa, and lack the unique heterokont signature sequence. They also appear not to be specifically related to either Amoebozoa or Radiolaria, with which they have sometimes been classified, so it is desirable to retain Heliozoa as a separate protozoan phylum. Even though centrohelids have no cilia or centrioles, the centrohelid clade branches among the bikont eukaryotes, but there is no strong bootstrap support for any particular position. Distance trees usually place centrohelids as sisters to haptophytes, whereas parsimony puts them as sisters to red algae, but there is no reason to think that either position is correct; both have very low bootstrap support. Quartet puzzling places them with fairly low support as sisters to the apusozoan zooflagellate Ancyromonas. As Ancyromonas is the only other eukaryote that shares the character combination of flat plate-like mitochondrial cristae and kinetocyst-type extrusomes with centrohelids, this position is biologically plausible, but because of weak support and conflict between trees it might not be correct. Irrespective of their precise position, our trees (together with previous evidence that Chlamydaster sterni has the derived dihydrofolate reductase/thymidylate synthetase gene fusion unique to bikonts) indicate that centrohelid heliozoa are ancestrally derived from a bikont flagellate by the loss of cilia. The centroplast that nucleates their axonemal microtubules is therefore almost certainly homologous with the centrosome of ciliated eukaryotes and should simply be called a centrosome.     

Ophiostomatoid fungi isolated from Japanese red pine and their relationships with bark beetles

We isolated ophiostomatoid fungi from bark beetles infesting Pinus densiflora and their galleries at 24 sites in Japan. Twenty-one ophiostomatoid fungi, including species of Ophiostoma, Grosmannia, Ceratocystiopsis, Leptographium, and Pesotum, were identified. Among these, 11 species were either newly recorded in Japan or were previously undescribed species. Some of these fungal species were isolated from several bark beetles, but other species were isolated from only a particular beetle species. Thus, it is suggested that some ophiostomatoid fungi have specific relationships with particular beetle species. In addition, fungus-beetle biplots from redundancy analysis (RDA) summarizing the effects of beetle ecological characteristics suggested that the association patterns between bark beetles and the associated fungi seemed to be related to the niches occupied by the beetles.     

Metabolites from Two Dominant Thermophilic Fungal Species Thermomyces lanuginosus and Scytalidium thermophilum

Thermomyces lanuginosus and Scytalidium thermophilum are among the most ubiquitous thermophilic fungi in compost and soil. Chemical study on these two prevalent strains collected from Yunnan led to isolation of 23 metabolites, including one new metabolite, therlanubutanolide, and 15 known compounds, isolated from the YGP culture broth of Thermomyces lanuginosus and 7 known compounds isolated from Scytalidium thermophilum, respectively. Therlanubutanolide shared the quite similar features of the same carbon skeleton and saturation as natural hexadecanoic acids. This was the first reported discovery of such a lactone as natural occurring metabolite. All the compounds were reported for the first time from thermophilic fungi. Among them, N‐[(2S,3R,4E,8E)‐1,3‐dihydroxy‐9‐methyloctadeca‐4,8‐dien‐2‐yl]acetamide was for the first time reported to be a naturally occurring metabolite and its NMR data was first provided in this study. A type of PKS‐derived metabolites, three 3,4‐dihydronaphthalen‐1(2H)‐ones, which were widely found in plant pathogenic fungi as phytotoxins and reported to have antimicrobial activity, were obtained from both dominant thermophilic fungi. The frequent occurrence of such PKS phytotoxins in these two thermophilic fungi might suggest particular ecological interest.     

Exosome isolation from hemolymph of Korean rhinoceros beetle,Allomyrina dichotoma (Coleoptera: Scarabaeidae)

Exosomes are 30–150 nm vesicles that are secreted from a range of cells. Recently, exosomes have been the subject of considerable research because there is mounting awareness of their diverse functions, including a role in cell–cell communication and presenting pathogens for immune responses. Exosomes contain diverse nucleic acid and protein cargos, derived not only from the organism but also from pathogens, making them suitable for use in disease diagnosis. The Korean rhinoceros beetle, Allomyrina dichotoma (Coleoptera: Scarabaeidae), is commercially reared in Korea for the pet trade and is used in traditional medicine for liver‐related diseases. However, several insect diseases caused by bacteria, fungi and viruses have been reported in A. dichotoma mass‐rearing facilities. Identifying these diseases with accuracy and in a timely manner is of paramount importance. Such diagnosis can be accomplished by identifying the nucleic acid or amino acid fragments from these disease‐causing pathogens in the exosome of A. dichotoma. We isolated exosomes from the hemolymph of A. dichotoma and used them to analyze exosome RNA and proteins. We confirmed the isolation of exosomes through RNA profiling, protein analysis and Western blotting. Our research established a solid foundation for using insect exosome protein and RNA analyses for the accurate diagnosis of insect diseases. To our knowledge, this is the first report of exosome isolation from insect hemolymph.     

The genetic basis for indole-diterpene chemical diversity in filamentous fungi

Indole-diterpenes are a structurally diverse group of secondary metabolites with a common cyclic diterpene backbone derived from geranylgeranyl diphosphate and an indole group derived from indole-3-glycerol phosphate. Different types and patterns of ring substitutions and ring stereochemistry generate this structural diversity. This group of compounds is best known for their neurotoxic effects in mammals, causing syndromes such as ‘ryegrass staggers’ in sheep and cattle. Because many of the fungi that synthesise these compounds form symbiotic relationships with plants, insects, and other fungi, the synthesis of these compounds may confer an ecological advantage to these associations. Considerable recent progress has been made on understanding indole-diterpene biosynthesis in filamentous fungi, principally through the cloning and characterisation of the genes and gene products for paxilline biosynthesis in Penicillium paxilli. Important insights into how the indole-diterpene backbone is synthesised and decorated have been obtained using P. paxilli mutants in this pathway. This review provides an overview of these recent developments.     

High selectivity in symbiotic associations of lichenized ascomycetes and cyanobacteria

The selectivity of mycobionts and cyanobionts in lichen symbioses were examined. We analyzed symbiotic cyanobionts, collected from different sample sites, and compared them to free‐living cyanobacteria Nostoc. Cyanobionts were obtained from lichens assigned to the genera Pseudocyphellaria and Sticta, in particular. Multiple gene loci were screened and direct optimization was used in the phylogenetic analyses. We show that many lichen fungi are strongly selective towards their cyanobionts. Lichenized ascomycetes seem to be able to identify and choose a specific strain, species or a species group of Nostoc with which to associate. The present analyses also suggest that some of the Nostoc taxa may be specialized in symbiotic life with only lichenized ascomycetes. Despite the selectivity observed in fungi, there appears to be no coevolution between the partners. We have also discussed the problems of using the tRNA^Leu intron as a marker in phylogenetic analyses. © The Willi Hennig Society 2006.     

Tryptophan‐derived secondary metabolites in Arabidopsis thaliana confer non‐host resistance to necrotrophic Plectosphaerella cucumerina fungi

A defence pathway contributing to non‐host resistance to biotrophic fungi in Arabidopsis involves the synthesis and targeted delivery of the tryptophan (trp)‐derived metabolites indol glucosinolates (IGs) and camalexin at pathogen contact sites. We have examined whether these metabolites are also rate‐limiting for colonization by necrotrophic fungi. Inoculation of Arabidopsis with adapted or non‐adapted isolates of the ascomycete Plectosphaerella cucumerina triggers the accumulation of trp‐derived metabolites. We found that their depletion in cyp79B2 cyp79B3 mutants renders Arabidopsis fully susceptible to each of three tested non‐adapted P. cucumerina isolates, and super‐susceptible to an adapted P. cucumerina isolate. This assigns a key role to trp‐derived secondary metabolites in limiting the growth of both non‐adapted and adapted necrotrophic fungi. However, 4‐methoxy‐indol‐3‐ylmethylglucosinolate, which is generated by the P450 monooxygenase CYP81F2, and hydrolyzed by PEN2 myrosinase, together with the antimicrobial camalexin play a minor role in restricting the growth of the non‐adapted necrotrophs. This contrasts with a major role of these two trp‐derived phytochemicals in limiting invasive growth of non‐adapted biotrophic powdery mildew fungi, thereby implying the existence of other unknown trp‐derived metabolites in resistance responses to non‐adapted necrotrophic P. cucumerina. Impaired defence to non‐adapted P. cucumerina, but not to the non‐adapted biotrophic fungus Erysiphe pisi, on cyp79B2 cyp79B3 plants is largely restored in the irx1 background, which shows a constitutive accumulation of antimicrobial peptides. Our findings imply differential contributions of antimicrobials in non‐host resistance to necrotrophic and biotrophic pathogens.     

Assessing the mycotoxigenic threat of necrotrophic pathogens of wheat

Pathogenic fungi are the causal agents of many significant plant diseases around the world. These diseases often result in significant yield reductions, leading to lower food production rates and economic losses. Several of these pathogenic fungi also produce mycotoxins during infection, which are harmful to human and animal health. Whilst some of these toxins and the fungi that produce them have been studied intensively, the mycotoxigenic potential of many of these pathogens remains largely unknown. Included within these fungi are the necrotrophic pathogens of wheat, Stagonospora nodorum, Pyrenophora tritici-repentis and Alternaria alternata. Recent studies have demonstrated that each of these pathogens is capable of synthesizing an array of mycotoxic compounds during disease development, questioning their status as non-mycotoxin producers. This review summarises recent mycotoxin findings in these necrotrophic wheat pathogens by briefly discussing the mycotoxins identified, their toxicity and their synthesis. Future and emerging threats are also considered.     

Presence in the atmosphere of vine and tomato pathogens

Summary In nature fungi, together with bacteria, play an important role in the degradation of organic substances. A small group of fungi is responsible for mycosis in man and animals. A larger group made up of different fungal taxa are particularly harmful to crops. The study of agricultural fungal phytopathologies is more difficult in that many taxa are characterized by a high degree of nutritional specialization. The identification in the field of the diseases caused by these taxa takes place often too late and not always are they controllable at a specific stage of their evolution. Since the majority of fungal spores is airborne and consequently present in various concentrations at different altitudes in the atmosphere, it is possible to monitor them and sometimes identify the fungus by sampling the air with a volumetric spore trap. Its diagnostic use has allowed to samplePlasmopara sporangia and obtain their concentration/m³ in the air of a vineyard. Similar results have been obtained in tomato fields, with the identification ofPhytophthora sporangia, the presence of which has been correlated to the meteorological data of the trial period. Field monitoring also offers the possibility of evaluating the presence of other fungi such as the generaBotrytis, Oidium, Alternaria, Pyricularia, Fusarium andStemphylium which damage crops.     

Hydroxy long-chain fatty acids in fungi

Hydroxy long-chain fatty acids occur widely in animals and plants and have important physiological activities in these eukaryotes. There are indications that these compounds are also common and important in fungi. The occurrence of hydroxy-polyunsaturated fatty acids (hydroxy-PUFAs) is of biotechnological importance, because these compounds are potentially high-value lipid products with medical applications. This review pays particular attention to the production of hydroxy-PUFAs by yeasts and other fungi. Hydroxy-PUFAs derived from lipoxygenase activity appear to be present in most fungi, while hydroxy-PUFAs from cyclooxygenase activity (i.e. prostaglandins) have mainly been implicated in the Oomycota and in yeasts from the genus Dipodascopsis. The occurrence of other hydroxy long-chain fatty acids in fungi is also discussed briefly; these include hydroxy fatty acids that are generally associated with cytochrome P-450 monooxygenase activity (i.e. terminal and sub-terminal hydroxy acids and diols derived from the corresponding epoxides) as well as 2-hydroxy-fatty acids and 3-hydroxy-fatty acids.The authors are with the Department of Microbiology and Biochemistry, University of the Orange Free State, P.O. Box 339, Bloemfontein, 9300, South Africa     

When to cut your losses: Dispersal allocation in an asexual filamentous fungus in response to competition

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Studies on inhibitory activities of antibodies against phytopathogenic fungi: an immunological approach

The major problem in agriculture is that of the fungal pathogens. In this era, where biological control is at focus, and is the centre of crop protection as well as environmental protection, the synthesis of new bio bodies is of utmost need. Fungicides available in the market, though of potential, are pathogen specific and highly pollutive. An attempt was made to raise polyclonal antibodies against Aspergillus niger. Following the particular standardised immunisation schedule, regular injections and periodic tapping were carried out. The IgG purified was used to check cross-reactivity with different crop fungi like Fusarium solani, Fusarium oxysporum f. sp. cicer (FOC), Rhizoctonia bataticola, Botryodiplodia theobromae, Curvularia sp., Alternaria porri and also with Aspergillus niger by two different methods. Liquid test media and the radial growth inhibition test performed in solid media were used to check the inhibition of fungi and cross-reactivity. The results were subjected to analysis of variance (ANOVA) by using Tukey's test at the significance level of p < 0.05. The antibodies were active against all the fungi for more than 15 days except for A. niger in which from the seventh day onwards spore germination was observed. The probable role of antibodies to detect the common antigenic molecule that may be present in all the tested fungi and their role in inhibiting these fungi is discussed.     

Composition and Nutritional Characteristics of Fungi Consumed by Callimico goeldii in Pando, Bolivia

Though ≥22 species of Primates consume fungi, most do so at low rates, comprising <5% of their feeding time. Goeldi's monkeys (Callimico goeldii), spend up to 29% of their feeding time year-round consuming fungal sporocarps, the fruiting bodies of fungi. We provide comprehensive data on the nutritional characteristics of 4 species of fungi consumed by Callimico goeldii (Ascopolyporus polyporoides, Ascopolyporus polychrous, Auricularia auricula, and Auricularia delicata). The composition of the fungi is similar to that of other fungi: predominantly fiber (66.2–83.0% dry matter) with small amounts of sugar (2.0–5.6% dry matter) and crude fat (0.9–1.6% dry matter). Though the crude protein content is substantial (5.5–13.4% dry matter), much of the nitrogen in the fungi is not likely to be available to Callimico goeldii because it is associated with indigestible food components or is in nonprotein form. The mineral content of the fungi are within the normal range for fungi generally and the calcium-to-phosphorus ratio is low (0.07–0.25). Fungi appear to be a low-quality food resource for Callimico goeldii and may contribute to their relatively large home ranges and low population density compared to other Callitrichinae. Research on the ability of Callimico goeldii to digest fungi is needed to understand fully the nutritional value of fungi to them. We discuss adaptations Callimico goeldii may have for improving their ability to obtain nutrients from fungi and potential ecological correlates of mycophagy.     

A quantitative structure–activity relationship study on some aromatic/heterocyclic sulfonamides and their charged derivatives acting as carbonic anhydrase inhibitors

A quantitative structure–activity relationship (QSAR) study is made on a series of aromatic/heterocyclic sulfonamides and their charged derivatives acting as carbonic anhydrase (CA) inhibitors. These compounds were studied by Scozzafava et al. (J. Med. Chem. 2000; 43: 292) for the selective inhibition of CAs—sulfonamides generally do not discriminate between different CA isozymes and hence exhibit many undesirable side effects when used as drugs against a particular disease. In this communication, an attempt has been made to investigate the physicochemical and structural properties that can make them selective for a given CA isozyme. Based on in vitro data reported by Scozzafava et al. against two cytosolic isozymes and one membrane-bound isozyme, the QSAR study has shown that uncharged compounds cannot be made selective for cytosolic or membrane-bound isozyme since in both the cases the compounds appear to follow the same mechanism of inhibition. However, for the charged compounds the polarizability of the molecule seems to greatly favor the inhibition of the membrane-bound enzyme, and hence they can be made selective for this enzyme by enhancing their polarizability, which is found to play no role in the inhibition of cytosolic enzymes.     

Primary simple assays of cellulose-degrading fungi

Some 25 fungi, including at least 14 basidiomycetes, one ascomycete, and five anamorphic fungi were evaluated for their cellulose-degrading abilities in Difco potato dextrose broth or Difco malt extract broth cultures with cellulosic substrates (e.g., filter paper) in plastic Petri dishes. Among them, Peniophora sp. 06-13 and Phlebia sp. 99-335 reduced the dry weights of the whole cultures with these substrates more than the dry weights of the respective original substrates after 30 days of culture, showing definite cellulose degradation. In the cultures with more than 10 test fungi including Pycnoporus coccineus 84-117, such weight losses did not occur. This assay technique for the primary screening for cellulose degrading fungi is simple, inexpensive, reproducible and accurate.     

Population‐specific effect of Wolbachia on the cost of fungal infection in spider mites

Many studies have revealed the ability of the endosymbiotic bacterium Wolbachia to protect its arthropod hosts against diverse pathogens. However, as Wolbachia may also increase the susceptibility of its host to infection, predicting the outcome of a particular Wolbachia‐host–pathogen interaction remains elusive. Yet, understanding such interactions and their eco‐evolutionary consequences is crucial for disease and pest control strategies. Moreover, how natural Wolbachia infections affect artificially introduced pathogens for biocontrol has never been studied. Tetranychus urticae spider mites are herbivorous crop pests, causing severe damage on numerous economically important crops. Due to the rapid evolution of pesticide resistance, biological control strategies using entomopathogenic fungi are being developed. However, although spider mites are infected with various Wolbachia strains worldwide, whether this endosymbiont protects them from fungi is as yet unknown. Here, we compared the survival of two populations, treated with antibiotics or naturally harboring different Wolbachia strains, after exposure to the fungal biocontrol agents Metarhizium brunneum and Beauveria bassiana. To control for potential effects of the bacterial community of spider mites, we also compared the susceptibility of two populations naturally uninfected by Wolbachia, treated with antibiotics or not. In one population, Wolbachia‐infected mites had a better survival than uninfected ones in absence of fungi but not in their presence, whereas in the other population Wolbachia increased the mortality induced by B. bassiana. In one naturally Wolbachia‐uninfected population, the antibiotic treatment increased the susceptibility of spider mites to M. brunneum, but it had no effect in the other treatments. These results suggest that natural Wolbachia infections may not hamper and may even improve the success of biological control using entomopathogenic fungi. However, they also draw caution on the generalization of such effects, given the complexity of within‐host–pathogens interaction and the potential eco‐evolutionary consequences of the use of biocontrol agents for Wolbachia‐host associations.     

The significance of peroxisomes in secondary metabolite biosynthesis in filamentous fungi

Peroxisomes are ubiquitous organelles characterized by a protein-rich matrix surrounded by a single membrane. In filamentous fungi, peroxisomes are crucial for the primary metabolism of several unusual carbon sources used for growth (e.g. fatty acids), but increasing evidence is presented that emphasize the crucial role of these organelles in the formation of a variety of secondary metabolites. In filamentous fungi, peroxisomes also play a role in development and differentiation whereas specialized peroxisomes, the Woronin bodies, play a structural role in plugging septal pores. The biogenesis of peroxisomes in filamentous fungi involves the function of conserved PEX genes, as well as genes that are unique for these organisms. Peroxisomes are also subject to autophagic degradation, a process that involves ATG genes. The interplay between organelle biogenesis and degradation may serve a quality control function, thereby allowing a continuous rejuvenation of the organelle population in the cells.     

A new function of trehalose and the peculiarities of lipid formation in mycelial fungi

This work deals with lipid formation in ascomycete fungi and the effect of preservatives on them. A new biological function of trehalose was revealed, and of particular interest was the fact that the effect of this disaccharide depended on its concentration in the growth medium. In the presence of a preservative such as potassium sorbate (PS), low trehalose concentrations suppressed the growth of mycelial fungi contaminating hard cheeses and contributed to the prolongation of the preservative’s effect. A tenfold increase in trehalose concentration in the medium, conversely, resulted in a drastic increase in growth activity and removed the PS effect. Therefore, trehalose can function as an inhibitor or a stimulator of growth processes, depending on its concentration. It was established that the secondary growth of Penicillium fungi during their ontogeny is accompanied by consumption of accumulated reserve lipids. In contrast, this phenomenon does not occur in mucorous fungi, and this probably accounts for the fact that Mucorales representatives can accumulate significant triacylglyceride amounts during the idiophase.     

VALVE AND BAND MORPHOLOGY OF SOME FRESHWATER DIATOMS. IV. OUTER SURFACE MUCILAGE OF NAVICULA CONFERVACEA VAR. CONFERVACEA1

The development of the mucilage on the outer surface of Navicula confervacea (Kütz.) Grun., a raphed, filamentous diatom, was studied with scanning electron microscopy. This nonstructural cell wall material, present on the surface after critical-point drying and absent after acid cleaning, was of two types: strands and papillae. Strands were associated with the raphe system, areolae, elongated pores of the mantle, and all girdle sutures. Organic papillae were a common feature of valves, valvo-copulae and pleurae, but their origin and distribution could not be explained since they often occurred between the obvious openings in the frustule. Strands from the raphe and areolae may function in attaching terminal cells to a substrate and adjacent cells to each other. Other strands of the girdle arise from sutures during cell enlargement and continue to lengthen and intertwine until the individual frustules within a filament are obscured. Strands from sutures might originate from the advalvar row of pores of the girdle bands since these pores lie along the suture, but direct observation of this was not made. Secretion between, the bands also cannot be ruled out. Although mucilaginous papillae may sometimes occur at random on the entire surface of frustules, there is also a distinct, narrow multiseriate row of them around the edge of valves without marginal spines.     

A review on the morphology of ovarian follicles in elasmobranchs: A case study in Rhizoprionodon taylori

The identification of the elasmobranch secondary ovarian follicles and their function can be challenging and the inconsistent use of terminology derived from other taxa is a matter of ongoing debate. In this study, the available information on the histology of the elasmobranch secondary ovarian follicles derived from atresia (preovulatory follicles) or ovulation (postovulatory follicles) is reviewed highlighting their morphology and steroidogenic capacity. Based on this literature review, the ovarian follicles of the Australian sharpnose shark Rhizoprionodon taylori were classified according to their preovulatory or postovulatory origin. Two types of secondary follicles originating from atresia of developing follicles (atretic previtellogenic follicles) and ripe follicles (atretic vitellogenic follicles), and one type of postovulatory follicle were identified throughout the reproductive year of this species. Morphological similarities of the elasmobranch secondary ovarian follicles and their variations in different species denote the difficulty to classify them. Given the multiple origins of ovarian follicles, their poorly understood functions and capacity to supply steroids, visual identification of these structures and the use of terminology derived from mammalian and other vertebrate studies (with the exception of the corpora lutea as a temporary endocrine gland) is not advisable. J. Morphol. 278:486–499, 2017. © 2017 Wiley Periodicals, Inc.