type histone deacetylase in the phytopathogenic
fungi, Magnaporthe oryzae (Pyricularia oryzae)
and Fusarium asiaticum. Lett Appl Microbiol
65:446–452
Margolin BS, Garrett-Engele PW, Stevens JN et al
(1998) A methylated Neurospora 5S rRNA pseudogene contains a transposable element inactivated by repeat-induced point mutation. Genetics
149:1787–1797
Martienssen R, Moazed D (2015) RNAi and heterochromatin assembly. Cold Spring Harb Perspect Biol 7:
a019323
Miao VP, Freitag M, Selker EU (2000) Short TpA-rich
segments of the zeta-eta region induce DNA methylation in Neurospora crassa. J Mol Biol 300:249–
273
Mo ¨ller M, Schotanus K, Soyer JL et al (2019) Destabilization of chromosome structure by histone H3
lysine 27 methylation. PLoS Genet 15:e1008093
Neigeborn L, Carlson M (1984) Genes affecting the
regulation of SUC2 gene expression by glucose
repression in Saccharomyces cerevisiae. Genetics
108:845–858
Niehaus E-M, Studt L, von Bargen KW et al (2016)
Sound of silence: the beauvericin cluster in Fusarium fujikuroi is controlled by cluster-specific and
global regulators mediated by H3K27 modification. Environ Microbiol 18:4282–4302
Nielsen PR, Nietlispach D, Mott HR et al (2002) Structure of the HP1 chromodomain bound to histone
H3 methylated at lysine 9. Nature 416:103–107
Olins AL, Olins DE (1974) Spheroid chromatin units (v
bodies). Science 183:330–332
Oudet P, Gross-Bellard M, Chambon P (1975) Electron
microscopic and biochemical evidence that chromatin structure is a repeating unit. Cell 4:281–300
Palmer JM, Bok JW, Lee S et al (2013) Loss of CclA,
required for histone 3 lysine 4 methylation,
decreases growth but increases secondary metabolite production in Aspergillus fumigatus. PeerJ 1:
e4
Pfannenstiel BT, Greco C, Sukowaty AT, Keller NP
(2018) The epigenetic reader SntB regulates secondary metabolism, development and global histone modifications in Aspergillus flavus. Fungal
Genet Biol 120:9–18
Pokholok DK, Harbison CT, Levine S et al (2005)
Genome-wide map of nucleosome acetylation and
methylation in yeast. Cell 122:517–527
Pueschel R, Coraggio F, Meister P (2016) From single
genes to entire genomes: the search for a function
of nuclear organization. Development 143:910–923
Rabl C (1885) U ¨ ber Zelltheilung. Morphologisches
Jahrbuch 10
Raduwan H, Isola AL, Belden WJ (2013) Methylation of
histone H3 on lysine 4 by the lysine methyltransferase SET1 protein is needed for normal clock
gene expression. J Biol Chem 288:8380–8390
Rea S, Eisenhaber F, O’Carroll D et al (2000) Regulation
of chromatin structure by site-specific histone H3
methyltransferases. Nature 406:593–599
Rountree MR, Selker EU (1997) DNA methylation
inhibits elongation but not initiation of transcription in Neurospora crassa. Genes Dev
11:2383–2395
Sancar C, Ha N, Yilmaz R et al (2015) Combinatorial
control of light induced chromatin remodeling
and gene activation in Neurospora. PLoS Genet
11:e1005105
Sasaki T, Lynch KL, Mueller CV et al (2014) Heterochromatin controls gH2A localization in Neurospora crassa. Eukaryot Cell 13:990–1000
Schmitt AD, Hu M, Ren B (2016) Genome-wide
mapping and analysis of chromosome architecture. Nat Rev Mol Cell Biol 17:743–755
Segal E, Widom J (2009) Poly(dA:dT) tracts: major
determinants of nucleosome organization. Curr
Opin Struct Biol 19:65–71
Selker EU (2002) Repeat-induced gene silencing in
fungi. Adv Genet 46:439–450
Selker EU, Cambareri EB, Jensen BC, Haack KR (1987)
Rearrangement of duplicated DNA in specialized
cells of Neurospora. Cell 51:741–752
Selker EU, Fritz DY, Singer MJ (1993) Dense nonsymmetrical DNA methylation resulting from repeatinduced point mutation in Neurospora. Science
262:1724–1728
Seymour M, Ji L, Santos AM et al (2016) Histone H1
limits DNA methylation in Neurospora crassa. G3
6:1879–1889
Shatkin AJ, Tatum EL (1959) Electron microscopy of
Neurospora crassa mycelia. J Biophys Biochem
Cytol 6:423–426
Shwab EK, Bok JW, Tribus M et al (2007) Histone
deacetylase activity regulates chemical diversity
in Aspergillus. Eukaryot Cell 6:1656–1664
Simon JA, Kingston RE (2013) Occupying chromatin:
polycomb mechanisms for getting to genomic targets, stopping transcriptional traffic, and staying
put. Mol Cell 49:808–824
Smith KM, Dobosy JR, Reifsnyder JE et al (2010) H2Band H3-specific histone deacetylases are required
for DNA methylation in Neurospora crassa. Genetics 186:1207–1216
Smith KM, Kothe GO, Matsen CB et al (2008) The
fungus Neurospora crassa displays telomeric
silencing mediated by multiple sirtuins and by
methylation of histone H3 lysine 9. Epigenetics
Chromatin 1:1–20
Smith KM, Phatale PA, Sullivan CM et al (2011) Heterochromatin is required for normal distribution
of Neurospora crassa CenH3. Mol Cell Biol
31:2528–2542
Solovei I, Thanisch K, Feodorova Y (2016) How to rule
the nucleus: divide et impera. Curr Opin Cell Biol
40:47–59
1 Chromatin Structure and Function in Neurospora crassa
23
fungi, Magnaporthe oryzae (Pyricularia oryzae)
and Fusarium asiaticum. Lett Appl Microbiol
65:446–452
Margolin BS, Garrett-Engele PW, Stevens JN et al
(1998) A methylated Neurospora 5S rRNA pseudogene contains a transposable element inactivated by repeat-induced point mutation. Genetics
149:1787–1797
Martienssen R, Moazed D (2015) RNAi and heterochromatin assembly. Cold Spring Harb Perspect Biol 7:
a019323
Miao VP, Freitag M, Selker EU (2000) Short TpA-rich
segments of the zeta-eta region induce DNA methylation in Neurospora crassa. J Mol Biol 300:249–
273
Mo ¨ller M, Schotanus K, Soyer JL et al (2019) Destabilization of chromosome structure by histone H3
lysine 27 methylation. PLoS Genet 15:e1008093
Neigeborn L, Carlson M (1984) Genes affecting the
regulation of SUC2 gene expression by glucose
repression in Saccharomyces cerevisiae. Genetics
108:845–858
Niehaus E-M, Studt L, von Bargen KW et al (2016)
Sound of silence: the beauvericin cluster in Fusarium fujikuroi is controlled by cluster-specific and
global regulators mediated by H3K27 modification. Environ Microbiol 18:4282–4302
Nielsen PR, Nietlispach D, Mott HR et al (2002) Structure of the HP1 chromodomain bound to histone
H3 methylated at lysine 9. Nature 416:103–107
Olins AL, Olins DE (1974) Spheroid chromatin units (v
bodies). Science 183:330–332
Oudet P, Gross-Bellard M, Chambon P (1975) Electron
microscopic and biochemical evidence that chromatin structure is a repeating unit. Cell 4:281–300
Palmer JM, Bok JW, Lee S et al (2013) Loss of CclA,
required for histone 3 lysine 4 methylation,
decreases growth but increases secondary metabolite production in Aspergillus fumigatus. PeerJ 1:
e4
Pfannenstiel BT, Greco C, Sukowaty AT, Keller NP
(2018) The epigenetic reader SntB regulates secondary metabolism, development and global histone modifications in Aspergillus flavus. Fungal
Genet Biol 120:9–18
Pokholok DK, Harbison CT, Levine S et al (2005)
Genome-wide map of nucleosome acetylation and
methylation in yeast. Cell 122:517–527
Pueschel R, Coraggio F, Meister P (2016) From single
genes to entire genomes: the search for a function
of nuclear organization. Development 143:910–923
Rabl C (1885) U ¨ ber Zelltheilung. Morphologisches
Jahrbuch 10
Raduwan H, Isola AL, Belden WJ (2013) Methylation of
histone H3 on lysine 4 by the lysine methyltransferase SET1 protein is needed for normal clock
gene expression. J Biol Chem 288:8380–8390
Rea S, Eisenhaber F, O’Carroll D et al (2000) Regulation
of chromatin structure by site-specific histone H3
methyltransferases. Nature 406:593–599
Rountree MR, Selker EU (1997) DNA methylation
inhibits elongation but not initiation of transcription in Neurospora crassa. Genes Dev
11:2383–2395
Sancar C, Ha N, Yilmaz R et al (2015) Combinatorial
control of light induced chromatin remodeling
and gene activation in Neurospora. PLoS Genet
11:e1005105
Sasaki T, Lynch KL, Mueller CV et al (2014) Heterochromatin controls gH2A localization in Neurospora crassa. Eukaryot Cell 13:990–1000
Schmitt AD, Hu M, Ren B (2016) Genome-wide
mapping and analysis of chromosome architecture. Nat Rev Mol Cell Biol 17:743–755
Segal E, Widom J (2009) Poly(dA:dT) tracts: major
determinants of nucleosome organization. Curr
Opin Struct Biol 19:65–71
Selker EU (2002) Repeat-induced gene silencing in
fungi. Adv Genet 46:439–450
Selker EU, Cambareri EB, Jensen BC, Haack KR (1987)
Rearrangement of duplicated DNA in specialized
cells of Neurospora. Cell 51:741–752
Selker EU, Fritz DY, Singer MJ (1993) Dense nonsymmetrical DNA methylation resulting from repeatinduced point mutation in Neurospora. Science
262:1724–1728
Seymour M, Ji L, Santos AM et al (2016) Histone H1
limits DNA methylation in Neurospora crassa. G3
6:1879–1889
Shatkin AJ, Tatum EL (1959) Electron microscopy of
Neurospora crassa mycelia. J Biophys Biochem
Cytol 6:423–426
Shwab EK, Bok JW, Tribus M et al (2007) Histone
deacetylase activity regulates chemical diversity
in Aspergillus. Eukaryot Cell 6:1656–1664
Simon JA, Kingston RE (2013) Occupying chromatin:
polycomb mechanisms for getting to genomic targets, stopping transcriptional traffic, and staying
put. Mol Cell 49:808–824
Smith KM, Dobosy JR, Reifsnyder JE et al (2010) H2Band H3-specific histone deacetylases are required
for DNA methylation in Neurospora crassa. Genetics 186:1207–1216
Smith KM, Kothe GO, Matsen CB et al (2008) The
fungus Neurospora crassa displays telomeric
silencing mediated by multiple sirtuins and by
methylation of histone H3 lysine 9. Epigenetics
Chromatin 1:1–20
Smith KM, Phatale PA, Sullivan CM et al (2011) Heterochromatin is required for normal distribution
of Neurospora crassa CenH3. Mol Cell Biol
31:2528–2542
Solovei I, Thanisch K, Feodorova Y (2016) How to rule
the nucleus: divide et impera. Curr Opin Cell Biol
40:47–59
1 Chromatin Structure and Function in Neurospora crassa
23