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Eric U Selker - One of the best experts on this subject based on the ideXlab platform.
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regional control of histone h3 lysine 27 methylation in Neurospora
Proceedings of the National Academy of Sciences of the United States of America, 2013Co-Authors: Kirsty Jamieson, Michael R Rountree, Zachary A Lewis, Jason E Stajich, Eric U SelkerAbstract:Trimethylated lysine 27 on histone H3 (H3K27me3) is present in Drosophila, Arabidopsis, worms, and mammals, but is absent from yeasts that have been examined. We identified and analyzed H3K27me3 in the filamentous fungus Neurospora crassa and in other Neurospora species. H3K27me3 covers 6.8% of the N. crassa genome, encompassing 223 domains, including 774 genes, all of which are transcriptionally silent. N. crassa H3K27me3-marked genes are less conserved than unmarked genes and only ∼35% of genes marked by H3K27me3 in N. crassa are also H3K27me3-marked in Neurospora discreta and Neurospora tetrasperma. We found that three components of the Neurospora Polycomb repressive complex 2 (PRC2)—[Su-(var)3–9; E(z); Trithorax] (SET)-7, embryonic ectoderm development (EED), and SU(Z)12 (suppressor of zeste12)—are required for H3K27me3, whereas the fourth component, Neurospora protein 55 (an N. crassa homolog of p55/RbAp48), is critical for H3K27me3 only at subtelomeric domains. Loss of H3K27me3, caused by deletion of the gene encoding the catalytic PRC2 subunit, set-7, resulted in up-regulation of 130 genes, including genes in both H3K27me3-marked and unmarked regions.
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gfp as a tool to analyze the organization dynamics and function of nuclei and microtubules in Neurospora crassa
Fungal Genetics and Biology, 2004Co-Authors: Michael Freitag, Eric U Selker, Namboori B. Raju, Patrick C Hickey, Nick D ReadAbstract:We report the construction of a versatile GFP expression plasmid and demonstrate its utility in Neurospora crassa. To visualize nuclei and microtubules, we generated carboxy-terminal fusions of sgfp to Neurospora histone H1 (hH1) and β-tubulin (Bml). Strong expression of GFP fusion proteins was achieved with the inducible Neurospora ccg-1 promoter. Nuclear and microtubule organization and dynamics were observed in live vegetative hyphae, developing asci, and ascospores by conventional and confocal laser scanning fluorescence microscopy. Observations of GFP fusion proteins in live cells largely confirmed previous results obtained by examination of fixed cells with various microscopic techniques. H1-GFP revealed dynamic nuclear shapes. Microtubules were mostly aligned parallel to the growth axis in apical compartments but more randomly arranged in sub-apical compartments. Time-lapse imaging of β-tubulin-GFP in germinating macroconidia revealed polymerization and depolymerization of microtubules. In heterozygous crosses, H1-GFP and β-tubulin-GFP expression was silenced, presumably by meiotic silencing. H1-GFP was translated in the vicinity of hH1+-sgfp+ nuclei in the common cytoplasm of giant Banana ascospores, but it diffused into all nuclei, another illustration of the utility of GFP fusion proteins.
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the genome sequence of the filamentous fungus Neurospora crassa
Nature, 2003Co-Authors: James E Galagan, Eric U Selker, Katherine A Borkovich, Sarah E Calvo, Nick O Read, David B Jaffe, William Fitzhugh, Serge Smirnov, Seth Purcell, Bushra RehmanAbstract:Neurospora crassa is a central organism in the history of twentieth-century genetics, biochemistry and molecular biology. Here, we report a high-quality draft sequence of the N. crassa genome. The approximately 40-megabase genome encodes about 10,000 protein-coding genes—more than twice as many as in the fission yeast Schizosaccharomyces pombe and only about 25% fewer than in the fruitfly Drosophila melanogaster. Analysis of the gene set yields insights into unexpected aspects of Neurospora biology including the identification of genes potentially associated with red light photobiology, genes implicated in secondary metabolism, and important differences in Ca 21 signalling as compared with plants and animals. Neurospora possesses the widest array of genome defence mechanisms known for any eukaryotic organism, including a process unique to fungi called repeat-induced point mutation (RIP). Genome analysis suggests that RIP has had a profound impact on genome evolution, greatly slowing the creation of new genes through genomic duplication and resulting in a genome with an unusually low proportion of closely related genes.
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the methylated component of the Neurospora crassa genome
Nature, 2003Co-Authors: Eric U Selker, Nikolaos A Tountas, Sally H Cross, Brian S Margolin, Jonathan G Murphy, Adrian Bird, Michael FreitagAbstract:Cytosine methylation is common, but not ubiquitous, in eukaryotes. Mammals1 and the fungus Neurospora crassa2,3 have about 2–3% of cytosines methylated. In mammals, methylation is almost exclusively in the under-represented CpG dinucleotides, and most CpGs are methylated1 whereas in Neurospora, methylation is not preferentially in CpG dinucleotides and the bulk of the genome is unmethylated4. DNA methylation is essential in mammals5 but is dispensable in Neurospora3,6, making this simple eukaryote a favoured organism in which to study methylation. Recent studies indicate that DNA methylation in Neurospora depends on one DNA methyltransferase, DIM-2 (ref. 6), directed by a histone H3 methyltransferase, DIM-5 (ref. 7), but little is known about its cellular and evolutionary functions. As only four methylated sequences have been reported previously in N. crassa, we used methyl-binding-domain agarose chromatography8 to isolate the methylated component of the genome. DNA sequence analysis shows that the methylated component of the genome consists almost exclusively of relics of transposons that were subject to repeat-induced point mutation—a genome defence system that mutates duplicated sequences9.
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a cytosine methyltransferase homologue is essential for repeat induced point mutation in Neurospora crassa
Proceedings of the National Academy of Sciences of the United States of America, 2002Co-Authors: Michael Freitag, Rebecca L Williams, Gregory O Kothe, Eric U SelkerAbstract:During sexual development, Neurospora crassa inactivates genes in duplicated DNA segments by a hypermutation process, repeat-induced point mutation (RIP). RIP introduces C:G to T:A transition mutations and creates targets for subsequent DNA methylation in vegetative tissue. The mechanism of RIP and its relationship to DNA methylation are not fully understood. Mutations in DIM-2, a DNA methyltransferase (DMT) responsible for all known cytosine methylation in Neurospora, does not prevent RIP. We used RIP to disrupt a second putative DMT gene in the Neurospora genome and tested mutants for defects in DNA methylation and RIP. No effect on DNA methylation was detected in the tissues that could be assayed, but the mutants showed recessive defects in RIP. Duplications of the am and mtr genes were completely stable in crosses homozygous for the mutated potential DMT gene, which we call rid (RIP defective). The same duplications were inactivated normally in heterozygous crosses. Disruption of the rid gene did not noticeably affect fertility, growth, or development. In contrast, crosses homozygous for a mutation in a related gene in Ascobolus immersus, masc1, reportedly fail to develop and heterozygous crosses reduce methylation induced premeiotically [Malagnac, F., Wendel, B., Goyon, C., Faugeron, G., Zickler, D., et al. (1997) Cell 91, 281–290]. We isolated homologues of rid from Neurospora tetrasperma and Neurospora intermedia to identify conserved regions. Homologues possess all motifs characteristic of eukaryotic DMTs and have large distinctive C- and N-terminal domains.
Michael Freitag - One of the best experts on this subject based on the ideXlab platform.
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gfp as a tool to analyze the organization dynamics and function of nuclei and microtubules in Neurospora crassa
Fungal Genetics and Biology, 2004Co-Authors: Michael Freitag, Eric U Selker, Namboori B. Raju, Patrick C Hickey, Nick D ReadAbstract:We report the construction of a versatile GFP expression plasmid and demonstrate its utility in Neurospora crassa. To visualize nuclei and microtubules, we generated carboxy-terminal fusions of sgfp to Neurospora histone H1 (hH1) and β-tubulin (Bml). Strong expression of GFP fusion proteins was achieved with the inducible Neurospora ccg-1 promoter. Nuclear and microtubule organization and dynamics were observed in live vegetative hyphae, developing asci, and ascospores by conventional and confocal laser scanning fluorescence microscopy. Observations of GFP fusion proteins in live cells largely confirmed previous results obtained by examination of fixed cells with various microscopic techniques. H1-GFP revealed dynamic nuclear shapes. Microtubules were mostly aligned parallel to the growth axis in apical compartments but more randomly arranged in sub-apical compartments. Time-lapse imaging of β-tubulin-GFP in germinating macroconidia revealed polymerization and depolymerization of microtubules. In heterozygous crosses, H1-GFP and β-tubulin-GFP expression was silenced, presumably by meiotic silencing. H1-GFP was translated in the vicinity of hH1+-sgfp+ nuclei in the common cytoplasm of giant Banana ascospores, but it diffused into all nuclei, another illustration of the utility of GFP fusion proteins.
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the methylated component of the Neurospora crassa genome
Nature, 2003Co-Authors: Eric U Selker, Nikolaos A Tountas, Sally H Cross, Brian S Margolin, Jonathan G Murphy, Adrian Bird, Michael FreitagAbstract:Cytosine methylation is common, but not ubiquitous, in eukaryotes. Mammals1 and the fungus Neurospora crassa2,3 have about 2–3% of cytosines methylated. In mammals, methylation is almost exclusively in the under-represented CpG dinucleotides, and most CpGs are methylated1 whereas in Neurospora, methylation is not preferentially in CpG dinucleotides and the bulk of the genome is unmethylated4. DNA methylation is essential in mammals5 but is dispensable in Neurospora3,6, making this simple eukaryote a favoured organism in which to study methylation. Recent studies indicate that DNA methylation in Neurospora depends on one DNA methyltransferase, DIM-2 (ref. 6), directed by a histone H3 methyltransferase, DIM-5 (ref. 7), but little is known about its cellular and evolutionary functions. As only four methylated sequences have been reported previously in N. crassa, we used methyl-binding-domain agarose chromatography8 to isolate the methylated component of the genome. DNA sequence analysis shows that the methylated component of the genome consists almost exclusively of relics of transposons that were subject to repeat-induced point mutation—a genome defence system that mutates duplicated sequences9.
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a cytosine methyltransferase homologue is essential for repeat induced point mutation in Neurospora crassa
Proceedings of the National Academy of Sciences of the United States of America, 2002Co-Authors: Michael Freitag, Rebecca L Williams, Gregory O Kothe, Eric U SelkerAbstract:During sexual development, Neurospora crassa inactivates genes in duplicated DNA segments by a hypermutation process, repeat-induced point mutation (RIP). RIP introduces C:G to T:A transition mutations and creates targets for subsequent DNA methylation in vegetative tissue. The mechanism of RIP and its relationship to DNA methylation are not fully understood. Mutations in DIM-2, a DNA methyltransferase (DMT) responsible for all known cytosine methylation in Neurospora, does not prevent RIP. We used RIP to disrupt a second putative DMT gene in the Neurospora genome and tested mutants for defects in DNA methylation and RIP. No effect on DNA methylation was detected in the tissues that could be assayed, but the mutants showed recessive defects in RIP. Duplications of the am and mtr genes were completely stable in crosses homozygous for the mutated potential DMT gene, which we call rid (RIP defective). The same duplications were inactivated normally in heterozygous crosses. Disruption of the rid gene did not noticeably affect fertility, growth, or development. In contrast, crosses homozygous for a mutation in a related gene in Ascobolus immersus, masc1, reportedly fail to develop and heterozygous crosses reduce methylation induced premeiotically [Malagnac, F., Wendel, B., Goyon, C., Faugeron, G., Zickler, D., et al. (1997) Cell 91, 281–290]. We isolated homologues of rid from Neurospora tetrasperma and Neurospora intermedia to identify conserved regions. Homologues possess all motifs characteristic of eukaryotic DMTs and have large distinctive C- and N-terminal domains.
Jay C. Dunlap - One of the best experts on this subject based on the ideXlab platform.
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Dissecting the mechanisms of the clock in Neurospora.
Methods in Enzymology, 2014Co-Authors: Jennifer M. Hurley, Jennifer J Loros, Jay C. DunlapAbstract:The circadian clock exists to synchronize inner physiology with the external world, allowing life to anticipate and adapt to the continual changes that occur in an organism's environment. The clock architecture is highly conserved, present in almost all major branches of life. Within eukaryotes, the filamentous fungus Neurospora crassa has consistently been used as an excellent model organism to uncover the basic circadian physiology and molecular biology. The Neurospora model has elucidated our fundamental understanding of the clock as nested positive and negative feedback loop, regulated by transcriptional and posttranscriptional processes. This review will examine the basics of circadian rhythms in the model filamentous fungus N. crassa as well as highlight the output of the clock in Neurospora and the reasons that N. crassa has continued to be a strong model for the study of circadian rhythms. It will also synopsize classical and emerging methods in the study of the circadian clock.
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6 Photobiology and Circadian Clocks in Neurospora
Fungal Genomics, 2014Co-Authors: Kevin K. Fuller, Jennifer J Loros, Jennifer M. Hurley, Jay C. DunlapAbstract:Light sensing and circadian rhythmicity are two related processes that promote the adaptation of many fungal species to their environment. This chapter begins with a description of fungal photoreceptors and their distributions across fungal lineages. We then discuss in some detail the molecular mechanisms of the photoresponse in two well-studied model fungi, Neurospora crassa and Aspergillus nidulans, placing an emphasis on the important similarities and differences between the two species. This will lead to a description of circadian rhythmicity in fungi in general, with a particular emphasis on Neurospora crassa. We highlight the core mechanism in this organism as well as discuss the inputs and outputs that can affect the clock. We also note at the end some new developments in molecular genetics in Neurospora.
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physical interaction between vivid and white collar complex regulates photoadaptation in Neurospora
Proceedings of the National Academy of Sciences of the United States of America, 2010Co-Authors: Chen-hui Chen, Jay C. Dunlap, Bradley S Demay, Amy S Gladfelter, Jennifer J LorosAbstract:Photoadaptation, the ability to attenuate a light response on prolonged light exposure while remaining sensitive to escalating changes in light intensity, is essential for organisms to decipher time information appropriately, yet the underlying molecular mechanisms are poorly understood. In Neurospora crassa, VIVID (VVD), a small LOV domain containing blue-light photoreceptor protein, affects photoadaptation for most if not all light-responsive genes. We report that there is a physical interaction between VVD and the white collar complex (WCC), the primary blue-light photoreceptor and the transcription factor complex that initiates light-regulated transcriptional responses in Neurospora. Using two previously characterized VVD mutants, we show that the level of interaction is correlated with the level of WCC repression in constant light and that even light-insensitive VVD is sufficient partly to regulate photoadaptation in vivo. We provide evidence that a functional GFP-VVD fusion protein accumulates in the nucleus on light induction but that nuclear localization of VVD does not require light. Constitutively expressed VVD alone is sufficient to change the dynamics of photoadaptation. Thus, our results demonstrate a direct molecular connection between two of the most essential light signaling components in Neurospora, VVD and WCC, illuminating a previously uncharacterized process for light-sensitive eukaryotic cells.
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Neurospora illuminates fungal photoreception.
Fungal Genetics and Biology, 2010Co-Authors: Chen-hui Chen, Jay C. Dunlap, Jennifer J LorosAbstract:Light not only is indispensable as an energy source for life on earth but also serves as an essential environmental cue conveying the information of daily and seasonal time to organisms across different kingdoms. Although the molecular mechanisms underlying light responses are actively explored in various light-sensitive organisms, these studies are either hindered by the complexity of the systems or an incomplete familiarity with the light signaling components involved in the scheme. Therefore, study of a simple and well-characterized model system is desirable to expand our knowledge of basic properties underlying the regulation of biological light responses. This review will briefly introduce the basic light sensing machinery in Neurospora crassa, a filamentous fungus, and then focus on the most recent advances in employing Neurospora as a model to study light signaling cascades, photoadaptation, and circadian clock-modulated effects in eukaryotic cells. Also, we will summarize the functions of a number of putative photoreceptors in Neurospora, and discuss the implications of the study of Neurospora to the field of fungal photobiology and some challenges for future studies.
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enabling a community to dissect an organism overview of the Neurospora functional genomics project
Advances in Genetics, 2007Co-Authors: Jay C. Dunlap, Kevin Mccluskey, Louise N Glass, Katherine A Borkovich, Matthew R Henn, Gloria E Turner, Matthew S Sachs, Michael Plamann, James E Galagan, Bruce W BirrenAbstract:A consortium of investigators is engaged in a functional genomics project centered on the filamentous fungus Neurospora, with an eye to opening up the functional genomic analysis of all the filamentous fungi. The overall goal of the four interdependent projects in this effort is to acccomplish functional genomics, annotation, and expression analyses of Neurospora crassa, a filamentous fungus that is an established model for the assemblage of over 250,000 species of nonyeast fungi. Building from the completely sequenced 43-Mb Neurospora genome, Project 1 is pursuing the systematic disruption of genes through targeted gene replacements, phenotypic analysis of mutant strains, and their distribution to the scientific community at large. Project 2, through a primary focus in Annotation and Bioinformatics, has developed a platform for electronically capturing community feedback and data about the existing annotation, while building and maintaining a database to capture and display information about phenotypes. Oligonucleotide-based microarrays created in Project 3 are being used to collect baseline expression data for the nearly 11,000 distinguishable transcripts in Neurospora under various conditions of growth and development, and eventually to begin to analyze the global effects of loss of novel genes in strains created by Project 1. cDNA libraries generated in Project 4 document the overall complexity of expressed sequences in Neurospora, including alternative splicing alternative promoters and antisense transcripts. In addition, these studies have driven the assembly of an SNP map presently populated by nearly 300 markers that will greatly accelerate the positional cloning of genes.
Christian Heintzen - One of the best experts on this subject based on the ideXlab platform.
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the Neurospora crassa circadian clock
Advances in Genetics, 2007Co-Authors: Christian HeintzenAbstract:Abstract The filamentous fungus Neurospora crassa is one of a handful of model organisms that has proven tractable for dissecting the molecular basis of a eukaryotic circadian clock. Work on Neurospora and other eukaryotic and prokaryotic organisms has revealed that a limited set of clock genes and clock proteins are required for generating robust circadian rhythmicity. This molecular clockwork is tuned to the daily rhythms in the environment via light‐ and temperature‐sensitive pathways that adjust its periodicity and phase. The circadian clockwork in turn transduces temporal information to a large number of clock‐controlled genes that ultimately control circadian rhythms in physiology and behavior. In summarizing our current understanding of the molecular basis of the Neurospora circadian system, this chapter aims to elucidate the basic building blocks of model eukaryotic clocks as we understand them today.
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coiled coil domain mediated frq frq interaction is essential for its circadian clock function in Neurospora
The EMBO Journal, 2001Co-Authors: Ping Cheng, Yuhong Yang, Christian HeintzenAbstract:The frequency (frq) gene, the central component of the frq-based circadian negative feedback loop, regulates various aspects of the circadian clock in Neurospora. However, the biochemical function of its protein products, FRQ, is poorly understood. In this study, we demonstrated that the most conserved region of FRQ forms a coiled-coil domain. FRQ interacts with itself in vivo, and the deletion of the coiled-coil region results in loss of the interaction. Point mutations, which are designed to disrupt the coiled-coil structure, weaken or completely abolish the FRQ self-association and lead to the arrhythmicity of the overt rhythm. Mutations of the FRQ coiled-coil that inhibit self-association also prevent its interaction with two other key components of the Neurospora circadian clock, namely WC-1 and WC-2, the two PAS domain-containing transcription factors. Taken together, these data strongly suggest that the formation of the FRQ–FRQ and FRQ–WC complexes is essential for the function of the Neurospora clock.
Jennifer J Loros - One of the best experts on this subject based on the ideXlab platform.
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Dissecting the mechanisms of the clock in Neurospora.
Methods in Enzymology, 2014Co-Authors: Jennifer M. Hurley, Jennifer J Loros, Jay C. DunlapAbstract:The circadian clock exists to synchronize inner physiology with the external world, allowing life to anticipate and adapt to the continual changes that occur in an organism's environment. The clock architecture is highly conserved, present in almost all major branches of life. Within eukaryotes, the filamentous fungus Neurospora crassa has consistently been used as an excellent model organism to uncover the basic circadian physiology and molecular biology. The Neurospora model has elucidated our fundamental understanding of the clock as nested positive and negative feedback loop, regulated by transcriptional and posttranscriptional processes. This review will examine the basics of circadian rhythms in the model filamentous fungus N. crassa as well as highlight the output of the clock in Neurospora and the reasons that N. crassa has continued to be a strong model for the study of circadian rhythms. It will also synopsize classical and emerging methods in the study of the circadian clock.
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Neurospora crassa looking back and looking forward at a model microbe
American Journal of Botany, 2014Co-Authors: Christine M Roche, Jennifer J Loros, Kevin Mccluskey, Louise N GlassAbstract:Investigation of the red bread mold that contaminated French bakeries nearly two centuries ago has led to a wealth of discoveries that have impacted our understanding of genetic, biochemical, and molecular mechanisms in microbes, from Mendelian genetics and the gene–enzyme relationship to circadian rhythm and plant cell wall degradation. Early Neurospora research focused on elucidating mechanisms of genetic recombination and gene action and later progressed to addressing complex biological questions of eukaryotic microbes. Here we review the evolution of the fi lamentous fungus Neurospora as a model microbe over the past century. We discuss the origins of Neurospora as a model microbe, the immediate scientifi c impacts from work in this fi lamentous fungus, and how the introduction of other model organisms (i.e., Escherichia coli and Saccharomyces cerevisiae ) redirected the focus of Neurospora research. Neurospora has and continues to inform our understanding of a myriad of basic scientifi c concepts and now has the opportunity to forge into the applied biosciences and biotechnology.
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6 Photobiology and Circadian Clocks in Neurospora
Fungal Genomics, 2014Co-Authors: Kevin K. Fuller, Jennifer J Loros, Jennifer M. Hurley, Jay C. DunlapAbstract:Light sensing and circadian rhythmicity are two related processes that promote the adaptation of many fungal species to their environment. This chapter begins with a description of fungal photoreceptors and their distributions across fungal lineages. We then discuss in some detail the molecular mechanisms of the photoresponse in two well-studied model fungi, Neurospora crassa and Aspergillus nidulans, placing an emphasis on the important similarities and differences between the two species. This will lead to a description of circadian rhythmicity in fungi in general, with a particular emphasis on Neurospora crassa. We highlight the core mechanism in this organism as well as discuss the inputs and outputs that can affect the clock. We also note at the end some new developments in molecular genetics in Neurospora.
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physical interaction between vivid and white collar complex regulates photoadaptation in Neurospora
Proceedings of the National Academy of Sciences of the United States of America, 2010Co-Authors: Chen-hui Chen, Jay C. Dunlap, Bradley S Demay, Amy S Gladfelter, Jennifer J LorosAbstract:Photoadaptation, the ability to attenuate a light response on prolonged light exposure while remaining sensitive to escalating changes in light intensity, is essential for organisms to decipher time information appropriately, yet the underlying molecular mechanisms are poorly understood. In Neurospora crassa, VIVID (VVD), a small LOV domain containing blue-light photoreceptor protein, affects photoadaptation for most if not all light-responsive genes. We report that there is a physical interaction between VVD and the white collar complex (WCC), the primary blue-light photoreceptor and the transcription factor complex that initiates light-regulated transcriptional responses in Neurospora. Using two previously characterized VVD mutants, we show that the level of interaction is correlated with the level of WCC repression in constant light and that even light-insensitive VVD is sufficient partly to regulate photoadaptation in vivo. We provide evidence that a functional GFP-VVD fusion protein accumulates in the nucleus on light induction but that nuclear localization of VVD does not require light. Constitutively expressed VVD alone is sufficient to change the dynamics of photoadaptation. Thus, our results demonstrate a direct molecular connection between two of the most essential light signaling components in Neurospora, VVD and WCC, illuminating a previously uncharacterized process for light-sensitive eukaryotic cells.
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Neurospora illuminates fungal photoreception.
Fungal Genetics and Biology, 2010Co-Authors: Chen-hui Chen, Jay C. Dunlap, Jennifer J LorosAbstract:Light not only is indispensable as an energy source for life on earth but also serves as an essential environmental cue conveying the information of daily and seasonal time to organisms across different kingdoms. Although the molecular mechanisms underlying light responses are actively explored in various light-sensitive organisms, these studies are either hindered by the complexity of the systems or an incomplete familiarity with the light signaling components involved in the scheme. Therefore, study of a simple and well-characterized model system is desirable to expand our knowledge of basic properties underlying the regulation of biological light responses. This review will briefly introduce the basic light sensing machinery in Neurospora crassa, a filamentous fungus, and then focus on the most recent advances in employing Neurospora as a model to study light signaling cascades, photoadaptation, and circadian clock-modulated effects in eukaryotic cells. Also, we will summarize the functions of a number of putative photoreceptors in Neurospora, and discuss the implications of the study of Neurospora to the field of fungal photobiology and some challenges for future studies.