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PubMed2003

The genetic basis of cellular morphogenesis in the filamentous fungus Neurospora crassa.

Cellular polarity is a fundamental property of every cell. Due to their extremely fast growth rate (>/=1 microm/s) and their highly elongated form, filamentous fungi represent a prime example of polarized growth and are an attractive model for the analysis of fundamental mechanisms underlying cellular polarity. To identify the critical components that contribute to polarized growth, we developed a large-scale genetic screen for the isolation of conditional mutants defective in this process in the model fungus Neurospora crassa. Phenotypic analysis and complementation tests of ca. 950 mutants identified more than 100 complementation groups that define 21 distinct morphological classes. The phenotypes include polarity defects over the whole hypha, more specific defects localized to hyphal tips or subapical regions, and defects in branch formation and growth directionality. To begin converting this mutant collection into meaningful biological information, we identified the defective genes in 45 mutants covering all phenotypic classes. These genes encode novel proteins as well as proteins which 1) regulate the actin or microtubule cytoskeleton, 2) are kinases or components of signal transduction pathways, 3) are part of the secretory pathway, or 4) have functions in cell wall formation or membrane biosynthesis. These findings highlight the dynamic nature of a fungal hypha and establish a molecular model for studies of hyphal growth and polarity.

Cell Polarity↗
PubMed2003

Maximal polar growth potential depends on the polarisome component AgSpa2 in the filamentous fungus Ashbya gossypii.

We used actin staining and videomicroscopy to analyze the development from a spore to a young mycelium in the filamentous ascomycete Ashbya gossypii. The development starts with an initial isotropic growth phase followed by the emergence of germ tubes. The initial tip growth speed of 6-10 microm/h increases during early stages of development. This increase is transiently interrupted in response to the establishment of lateral branches or septa. The hyphal tip growth speed finally reaches a maximum of up to 200 micro/h, and the tips of these mature hyphae have the ability to split into two equally fast-growing hyphae. A search for A. gossypii homologs of polarisome components of the yeast Saccharomyces cerevisiae revealed a remarkable size difference between Spa2p of both organisms, with AgSpa2p being double as long as ScSpa2p due to an extended internal domain. AgSpa2 colocalizes with sites of polarized actin. Using time-lapse videomicroscopy, we show that AgSpa2p-GFP polarization is established at sites of branch initiation and then permanently maintained at hyphal tips. Polarization at sites of septation is transient. During apical branching the existing AgSpa2p-GFP polarization is symmetrically divided. To investigate the function of AgSpa2p, we generated two AgSPA2 mutants, a partial deletion of the internal domain alone, and a complete deletion. The mutations had an impact on the maximal hyphal tip growth speed, on the hyphal diameter, and on the branching pattern. We suggest that AgSpa2p is required for the determination of the area of growth at the hyphal tip and that the extended internal domain plays an important role in this process.

Actins↗
PubMed2004

A Ras-like GTPase is involved in hyphal growth guidance in the filamentous fungus Ashbya gossypii.

Characteristic features of morphogenesis in filamentous fungi are sustained polar growth at tips of hyphae and frequent initiation of novel growth sites (branches) along the extending hyphae. We have begun to study regulation of this process on the molecular level by using the model fungus Ashbya gossypii. We found that the A. gossypii Ras-like GTPase Rsr1p/Bud1p localizes to the tip region and that it is involved in apical polarization of the actin cytoskeleton, a determinant of growth direction. In the absence of RSR1/BUD1, hyphal growth was severely slowed down due to frequent phases of pausing of growth at the hyphal tip. During pausing events a hyphal tip marker, encoded by the polarisome component AgSPA2, disappeared from the tip as was shown by in vivo time-lapse fluorescence microscopy of green fluorescent protein-labeled AgSpa2p. Reoccurrence of AgSpa2p was required for the resumption of hyphal growth. In the Agrsr1/bud1Delta deletion mutant, resumption of growth occurred at the hyphal tip in a frequently uncoordinated manner to the previous axis of polarity. Additionally, hyphal filaments in the mutant developed aberrant branching sites by mislocalizing AgSpa2p thus distorting hyphal morphology. These results define AgRsr1p/Bud1p as a key regulator of hyphal growth guidance.

Actins↗
PubMed2002

YlBMH1 encodes a 14-3-3 protein that promotes filamentous growth in the dimorphic yeast Yarrowia lipolytica.

Most pathogenic fungi have the ability to alternate between a unicellular yeast form and different filamentous forms (hyphae and pseudohyphae). This attribute is generally regarded as an important virulence factor and has also attracted attention because of its implications in the study of eukaryotic cell differentiation. To identify genes that are involved in the regulation of these events, chemical mutagenesis of the dimorphic yeast Yarrowia lipolytica was performed and morphological mutants that were unable to form hyphal cells were isolated. Screening of a Y. lipolytica genomic DNA library for genes able to complement this defect led to the isolation of YlBMH1, a gene encoding a 14-3-3 protein and whose transcription levels are increased during the yeast-to-hypha transition. Remarkably, overexpression of YlBMH1 was able to enhance pseudohyphae formation in a strain lacking functional YlRAC1 but caused no visible effects in deltamhy1 and deltabem1 cells, thus suggesting that YlBMH1 is involved in the regulation of both hyphal and pseudohyphal growth in Y. lipolytica. The identification of YlBMH2, a gene encoding a second 14-3-3 protein (YlBmh2p) that contains a 19 aa insertion absent in all other members of the 14-3-3 family, is also reported. Differently from YlBMH1, the transcription levels of YlBMH2 do not show any apparent variation during the induction of hyphal growth, and its overexpression has no effects on cells lacking functional MHY1, YlRAC1 or YlBEM1. Taken together, these observations suggest that, in spite of their high conservation, YlBmh1p and YlBmh2p have different cellular functions.

14-3-3 Proteins↗
PubMed2002

Repression of chsB expression reveals the functional importance of class IV chitin synthase gene chsD in hyphal growth and conidiation of Aspergillus nidulans.

The functions of two previously identified chitin synthase genes in Aspergillus nidulans, chsB and chsD, were analysed. First, a conditional chsB mutant was constructed in which the expression of chsB is under the control of a repressible promoter, the alcA promoter, of A. nidulans. Under repressing conditions, the mutant grew slowly and produced highly branched hyphae, supporting the idea that chsB is involved in normal hyphal growth. The involvement of chsB in conidiation was also demonstrated. Next, double mutants of chsB and chsD were constructed, in which chsB was placed under the control of the alcA promoter and chsD was replaced with the argB gene of A. nidulans. These double mutants grew more slowly than the chsB single mutant under high-osmolarity conditions. The hyphae of the double mutant appeared to be more disorganized than those of the chsB single mutant. It was also found that ChsD was functionally implicated in conidiation when the expression of chsB was limited. These results indicate the importance of the ChsD function in the absence of chsB expression. The roles of ChsB and ChsD in hyphal growth and in conidiation were supported by the analysis of the spatial expression patterns of chsB and chsD, using lacZ of Escherichia coli as a reporter gene.

Aspergillus nidulans↗
PubMed2026

Depletion of the Candida albicans TLO gene family reveals a requirement for alpha TLO genes for wild-type virulence.

Candida albicans uniquely possesses an expanded family of genes (the TLO gene family) that encodes 10-15 paralogues of the Med2 component of the transcriptional regulator Mediator. Previous studies have shown that TLO null mutants are unable to form hyphae and are hypersensitive to environmental stress. However, the reason for the TLO gene expansion remains unclear, and the current study aimed to determine if reduction in the TLO family copy number affected virulence. In order to investigate this, we used CRISPR-Cas9 mutagenesis to generate two TLO-depleted mutants: one mutant retaining only TLOβ2 (CaTLO2) and the second mutant containing only TLOγ5 (CaTLO5). Both TLO-depleted mutants exhibited increased filamentous growth, increased susceptibility to specific stresses and reduced virulence in a murine model of oropharyngeal candidiasis (OPC). In vitro, the CaTLO5 mutant also exhibited impaired hyphal escape from macrophages and reduced hyphal invasion of oral keratinocytes. We then investigated if complementation with TLOα1, a gene previously shown to restore wild-type growth in a Δtlo null mutant, could restore virulence. In vitro infection models showed that TLOα1 could restore true hypha formation, epithelial invasion and hyphal escape from macrophages in the CaTLO5 background. The murine OPC model showed that TLOα1 could restore wild-type virulence in both CaTLO2 and CaTLO5 strains, suggesting an essential role for α-TLO in oral mucosal infection. Together, these findings highlight the functional specialization between the α, β and γ TLO gene groups and establish α-TLO as a major regulator of virulence in C. albicans.

Candida albicans↗
PubMed2004

The GPI-anchored protein CaEcm33p is required for cell wall integrity, morphogenesis and virulence in Candida albicans.

Ecm33p is a widely distributed fungal protein with functional relevance, clearly demonstrated by ecm33Delta mutant phenotypes, mainly related to the cell wall. Homology searches with Saccharomyces cerevisiae genes identified Candida albicans Ecm33p, as well as the two other proteins of its family: Pst1p and the product of YCL048w. C. albicans Ecm33p is a 423 aa protein which has the typical features of cell-surface GPI proteins and is able to complement S. cerevisiae ecm33Delta cell wall defects. Heterozygous (RML1) and homozygous (RML2) mutants of CaECM33 were obtained, as well as a single and a double reintegrant (RML3 and RML4, respectively). Caecm33 mutant strains displayed an aberrant morphology, being more rounded and bigger than the wild-type, suggesting morphogenetic defects. They also exhibited cell wall defects, with enhanced sensitivity to different compounds that interfere in polymerization of cell wall components (Calcofluor white, Congo red and hygromycin B) and a marked tendency to flocculate extensively. In addition, CaEcm33p is required for normal C. albicans yeast-to-hyphae transition in vitro. In liquid medium (5 % serum), the transition was delayed in Caecm33 mutants, and after 24 h the culture contained very abnormal large and rounded cells. On solid medium (10 % serum, Spider or SLADH) RML2 failed to produce hyphae and media invasiveness. CaECM33 showed a gene dosage effect, demonstrated by the intermediate phenotype of the heterozygous mutants RML1 and confirmed by Northern blot analysis. Furthermore, CaEcm33p is also involved in C. albicans virulence. In a murine systemic model of infection, 100 % mouse survival and no kidney or brain colonization were obtained 30 days after infection with 10(6) Candida cells of any homozygous or heterozygous Caecm33Delta mutant tested. In contrast, all mice infected with parental or RML4 (two CaECM33 copies reintegrated) strains died in a few days, showing that, in these conditions, two CaECM33 copies were required for virulence.

Animals↗
PubMed2006

Effects of amino acid on morphological development and nucleus formation of arachidonic acid-producing filamentous micro-organism, Mortierella alpina.

AIMS: Effects of amino acid on morphological development and nucleus formation of arachidonic acid-producing filamentous micro-organism, Mortierella alpina were investigated using flow-through chamber. METHODS AND RESULTS: Mortierella alpina CBS 754.68 was cultivated in flow through chamber using nutrient-rich, minimal and specific amino acid-containing minimal media. To investigate the effect of amino acid on morphological parameters either 0.28 g l(-1) alanine, 0.53 g l(-1) sodium glutamate one hydrate or 0.42 g l(-1) valine was added to the minimal medium. In a flow-through chamber, the growth of hyphal elements and nucleus formation of arachidonic acid-producing fungus M. alpina were studied on-line, using image analysis techniques. When the Ala- and Val-containing media were used, the hyphal growth units (HGUs) were 90.2 and 86.7 microm per tip, respectively, which were 2.4-fold higher than that in the nutrient-rich medium, indicating that Ala and Val stimulate the elongation of hyphae. The specific nucleus formation rates were Glu->Val-containing media>minimal and nutrient-rich media>Ala-containing medium. The nucleus doubling times in Glu- and Val-containing media were 1.9 and 2 h, respectively, which were not significant different. CONCLUSIONS: Ala and Val stimulate the elongation of M. alpina hyphae, and nucleus formation rates were Glu->Val->Ala-containing media. SIGNIFICANCE AND IMPACT OF THE STUDY: Formation of fungal morphology and nucleus were shown using the flow-through chamber coupled with image analysis, which making possible to discuss the relationship between mycelial morphology and nucleus formation of M. alpina.

Alanine↗
PubMed2004

Regulators of G-protein signalling in Aspergillus nidulans: RgsA downregulates stress response and stimulates asexual sporulation through attenuation of GanB (Galpha) signalling.

Regulators of G-protein signalling play a crucial role in controlling the degree of heterotrimeric G-protein signalling. In addition to the previously studied flbA, we have identified three genes (rgsA, rgsB and rgsC) encoding putative RGS proteins in the genome of Aspergillus nidulans. Characterization of the rgsA gene revealed that RgsA downregulates pigment production and conidial germination, but stimulates asexual sporulation (conidiation). Deletion of rgsA (DeltargsA) resulted in reduced colony size with increased aerial hyphae, elevated accumulation of brown pigments as well as enhanced tolerance of conidia and vegetative hyphae against oxidative and thermal stress. Moreover, DeltargsA resulted in conidial germination in the absence of a carbon source. Deletion of both flbA and rgsA resulted in an additive phenotype, suggesting that the G-protein pathways controlled by FlbA and RgsA are different. Morphological and metabolic alterations caused by DeltargsA were suppressed by deletion of ganB encoding a Galpha subunit, indicating that the primary role of RgsA is to control negatively GanB-mediated signalling. Overexpression of rgsA caused inappropriate conidiation in liquid submerged culture, supporting the idea that GanB signalling represses conidiation. Our findings define a second and specific RGS-Galpha pair in A. nidulans, which may govern upstream regulation of fungal cellular responses to environmental changes.

Adaptation, Physiological↗
PubMed2005

Relationship between Candida infection and immune cellular response in inflammatory hyperplasia.

OBJECTIVES: To analyze and quantify the CD8(+) and CD4(+) T-lymphocyte populations in inflammatory hyperplasia and to establish the relationship between the frequency and location of these cells and Candida infection. METHODS: Samples of inflammatory hyperplasia were stained with PAS for evidence of Candida sp. and were classified in two groups, infected and control, according to the presence or absence of infection. After immunoreaction with specific anti-CD4 and anti-CD8 monoclonal antibodies, the distribution and frequency of the positive cells were analyzed in 41 cases (19 controls without Candida sp. and 22 infected cases). Lymphocytes were quantified in the three consecutive fields where the inflammatory infiltration was concentrated. RESULTS: There was no relationship between the frequency and location of CD4(+) T cells and Candida sp. infection. The number of CD8(+) cells close to the fungi hyphae as well as the total number of CD8(+) T cells present in inflammatory hyperplasia were higher in the Candida sp. group than in the control noninfected group (P < 0.05). CONCLUSION: Since the CD8(+) T cells were distributed according to the location of Candida sp. hyphae, and since a higher CD8(+)/total lymphocytes ratio was observed in the infected group, we suggest a role for CD8(+) T cells in the defense against Candida in oral infections associated with inflammatory hyperplasia in immunocompetent individuals.

CD4-Positive T-Lymphocytes↗
PubMed2005

Arbuscular mycorrhizal fungi reveal distinct patterns of anastomosis formation and hyphal healing mechanisms between different phylogenic groups.

The significance of anastomosis formation and the hyphal healing mechanism (HHM) for functionality and integrity of the arbuscular mycorrhizal (AM) fungal mycelial network remains poorly documented. Four Glomeraceae and three Gigasporaceae were cultured monoxenically. Anastomosis formation was assessed using the grid line method, while HHM was time-lapse monitored. In intact mycelial networks, the number of anastomosis per hyphal length was higher for Glomeraceae than for Gigasporaceae strains. Glomeraceae strains studied always formed anastomosis between different hyphae, whereas anastomosis in the Gigasporaceae more often concerned hyphal bridges within the same hyphae. In both families the HHM corresponded to a four-step process; first septum formation; second initiation of growing hyphal tips (GHTs); third GHT elongation, orientation and contact; and fourth GHT fusion and cytoplasmic/protoplasmic flux re-establishment. These four steps differentiated Glomeraceae from Gigasporaceae. The type and number of anastomosis per hyphal length, and the HHM differed considerably between Glomeraceae and Gigasporaceae families representing a supplementary character that distinguishes these two families and may be of significance in ecological studies of AM fungi.

Cytoplasm↗
PubMed2006

Modelling the contribution of arbuscular mycorrhizal fungi to plant phosphate uptake.

In this paper, we investigate the role of arbuscular mycorrhizal fungi in plant phosphorus nutrition. We develop a mathematical model which quantitatively assesses the contribution of external fungal hyphae to plant phosphate uptake. We derive an equation for solute uptake by a growing fungal mycelium which we couple with a model for root uptake. We analyse the model using nondimensionalization and numerical simulations. Simulations predict that removal of phosphate from soil is dominated by hyphal uptake as opposed to root uptake. Model analysis shows that the depletion zones around hyphae overlap within 8 h and that the transfer between fungus and root is a critical step for the behaviour of phosphorus within the mycelial phase. We also show that the volume fraction of mycelium is negligibly small in comparison to other soil phases. This is the first model to quantify the contribution of mycorrhizal fungi to plant phosphate uptake. A full data set for model parametrization and validation is not currently available. Therefore, more complete sets of experimental measurements are necessary to make this model more applicable.

Hyphae↗
PubMedDate not supplied

Analysis of morphological relationship between micro- and macromorphology of Mortierella species using a flow-through chamber coupled with image analysis.

Using a flow-through chamber coupled with image analysis, the morphological parameters of 11 Mortierella species were quantified, and the relationship between micro- and macromorphology was investigated. On potato-dextrose-agar plates, 5 species formed rose petal-like colonies, 3 formed large round colonies, and 3 formed donut-like colonies. By observing micromorphology in a flow-through chamber, fungi were divided into 3 groups, classified according to morphological parameters: (i) a group with a high branch formation rate (q(b): tip/microm/h) and a low tip extension rate (q(tip): microm/tip/h); (ii) a group with a low branch formation rate and a high tip extension rate; and (iii) a group intermediate between the former and the latter groups. In suspension culture, group (i) fungi formed a hyphal bundle with a pulpy pellet-like morphology and a pellet core. In contrast, group (ii) fungi showed an aggregation of hyphae without the pellet core. In a narrow-specific hyphal growth rate (mu(l)) range (0.35-0.45 h(-1)), a higher branch formation rate led to increased hyphal branching, resulting in the formation of a hyphal bundle with a pulpy pellet-like morphology and a pellet core. When the branch formation rate was lower than 2 x 10(-3) tips/microm/h, the mycelia formed less branched but longer hypha. Our study surmises that a micromorphology consisting of a high hyphal growth rate (0.4 h(-1)), low tip extension rate (20 tips/microm/h), and high branch formation rate (8 x 10(-3) tips/microm/h) forms the suitable macromorphology for arachidonic acid production.

Hyphae↗
PubMed2002

Aspergillus oryzae in solid-state and submerged fermentations. Progress report on a multi-disciplinary project.

We report the progress of a multi-disciplinary research project on solid-state fermentation (SSF) of the filamentous fungus Aspergillus oryzae. The molecular and physiological aspects of the fungus in submerged fermentation (SmF) and SSF are compared and we observe a number of differences correlated with the different growth conditions. First, the aerial hyphae which occur only in SSFs are mainly responsible for oxygen uptake. Second, SSF is characterised by gradients in temperature, water activity and nutrient concentration, and inside the hyphae different polyols are accumulating. Third, pelleted growth in SmF and mycelial growth in SSF show different gene expression and protein secretion patterns. With this approach we aim to expand our knowledge of mechanisms of fungal growth on solid substrates and to exploit the biotechnological applications.

Aspergillus oryzae↗
PubMed2006

Functional analysis of the ATG8 homologue Aoatg8 and role of autophagy in differentiation and germination in Aspergillus oryzae.

Autophagy is a well-known degradation system, induced by nutrient starvation, in which cytoplasmic components and organelles are digested via vacuoles/lysosomes. Recently, it was reported that autophagy is involved in the turnover of cellular components, development, differentiation, immune responses, protection against pathogens, and cell death. In this study, we isolated the ATG8 gene homologue Aoatg8 from the filamentous fungus Aspergillus oryzae and visualized autophagy by the expression of DsRed2-AoAtg8 and enhanced green fluorescent protein-AoAtg8 fusion proteins in this fungus. While the fusion proteins were localized in dot structures which are preautophagosomal structure-like structures under normal growth conditions, starvation or rapamycin treatment caused their accumulation in vacuoles. DsRed2 expressed in the cytoplasm was also taken up into vacuoles under starvation conditions or during the differentiation of conidiophores and conidial germination. Deletion mutants of Aoatg8 did not form aerial hyphae and conidia, and DsRed2 was not localized in vacuoles under starvation conditions, indicating that Aoatg8 is essential for autophagy. Furthermore, Aoatg8 conditional mutants showed delayed conidial germination in the absence of nitrogen sources. These results suggest that autophagy functions in both the differentiation of aerial hyphae and in conidial germination in A. oryzae.

Aspergillus oryzae↗
PubMed2006

Tcc1p, a novel protein containing the tetratricopeptide repeat motif, interacts with Tup1p to regulate morphological transition and virulence in Candida albicans.

The transcriptional factor CaTup1p represses many genes involved in intracellular processes, including the yeast-hypha transition, in the human fungal pathogen Candida albicans. Using tandem affinity purification technology, we identified a novel protein that interacts with CaTup1p, named Tcc1p (Tup1p complex component). Tcc1p is a C. albicans-specific protein with a 736-amino-acid polypeptide with four tetratricopeptide repeat (TPR) motifs in the N-terminal portion. Tcc1p formed a protein complex with CaTup1p via the TPR domain of Tcc1p, independently of CaSsn6p-CaTup1p The tcc1Delta disruptant showed filamentous growth under conditions inducing the yeast form, as is true of the Catup1Delta mutant. Consistent with this result, the common set of hypha-specific genes was negatively regulated by both TCC1 and CaTUP1. These observations will provide new insights into CaTup1p-dependent transcriptional gene regulation in C. albicans.

Amino Acid Sequence↗
PubMed2004

Lipid raft polarization contributes to hyphal growth in Candida albicans.

The polarization of sterol- and sphingolipid-enriched domains (lipid rafts) has been linked to morphogenesis and cell movement in diverse cell types. In the yeast Saccharomyces cerevisiae, a dramatic polarization of sterol-rich domains to the shmoo tip was observed in pheromone-induced cells (M. Bagnat and K. Simons, Proc. Natl. Acad. Sci. USA 99:14183-14188, 2002). We therefore examined whether plasma membrane lipid polarization contributes to the ability of the fungal pathogen Candida albicans to grow in a highly polarized manner to form hyphae. Interestingly, staining with filipin revealed that membrane sterols were highly polarized to the leading edge of growth during all stages of hyphal growth. Budding and pseudohyphal cells did not display polarized staining. Filipin staining was also enriched at septation sites in hyphae, where colocalization with septin proteins was observed, suggesting a role for the septins in forming a boundary domain. Actin appeared to play a role in sterol polarization and hyphal morphogenesis in that both were disrupted by low concentrations of latrunculin A that did not prevent budding. Furthermore, blocking either sphingolipid biosynthesis with myriocin or sterol biosynthesis with ketoconazole resulted in a loss of ergosterol polarization and caused abnormal hyphal morphogenesis, suggesting that lipid rafts are involved. Since hyphal growth is required for the full virulence of C. albicans, these results suggest that membrane polarization may contribute to the pathogenesis of this organism.

Antifungal Agents↗
PubMed2005

Release from quorum-sensing molecules triggers hyphal formation during Candida albicans resumption of growth.

Candida albicans is a pathogenic fungus able to change morphology in response to variations in its growth environment. Simple inoculation of stationary cells into fresh medium at 37 degrees C, without any other manipulations, appears to be a powerful but transient inducer of hyphal formation; this process also plays a significant role in classical serum induction of hyphal formation. The mechanism appears to involve the release of hyphal repression caused by quorum-sensing molecules in the growth medium of stationary-phase cells, and farnesol has a strong but incomplete role in this process. We used DNA microarray technology to study both the resumption of growth of Candida albicans cells and molecular regulation involving farnesol. Maintaining farnesol in the culture medium during the resumption of growth both delays and reduces the induction of hypha-related genes yet triggers expression of genes encoding drug efflux components. The persistence of farnesol also prevents the repression of histone genes during hyphal growth and affects the expression of putative or demonstrated morphogenesis-regulating cyclin genes, such as HGC1, CLN3, and PCL2. The results suggest a model explaining the triggering of hyphae in the host based on quorum-sensing molecules.

Candida albicans↗