quantitative-live-cell-fluorescence-microscopy-analysis-fission

Quantitative Live Cell Fluorescence-microscopy Analysis of Fission Yeast

Several microscopy techniques are available today that can detect a specific protein within the cell. During the last decade live cell imaging using ...

Histone deacetylases (HDACs) are important for gene regulation and the maintenance of heterochromatin in eukaryotes. Schizosaccharomyces pombe was used as a model system to investigate the functional divergence within this conserved enzyme family. S. pombe has three HDACs encoded by the hda1(+), clr3(+), and clr6(+) genes. Strains mutated in these genes have previously been shown to display strikingly different phenotypes when assayed for viability, chromosome loss, and silencing. Here, conserved differences in the substrate binding pocket identify Clr6 and Hda1 as class I HDACs, while Clr3 belongs in the class II family. Furthermore, these HDACs were shown to have strikingly different subcellular localization patterns. Hda1 was localized to the cytoplasm, while most of Clr3 resided throughout the nucleus. Finally, Clr6 was localized exclusively on the chromosomes in a spotted pattern. Interestingly, Clr3, the only HDAC present in the nucleolus, was required for ribosomal DNA (rDNA) silencing. Clr3 presumably acts directly on heterochromatin, since it colocalized with the centromere, mating-type region, and rDNA as visualized by in situ hybridization. In addition, Clr3 could be cross-linked to mat3 in chromatin immunoprecipitation experiments. Western analysis of bulk histone preparations indicated that Hda1 (class I) had a generally low level of activity in vivo and Clr6 (class I) had a high level of activity and broad in vivo substrate specificity, whereas Clr3 (class II) displayed its main activity on acetylated lysine 14 of histone H3. Thus, the distinct functions of the S. pombe HDACs are likely explained by their distinct cellular localization and their different in vivo specificities.

Functional divergence between histone deacetylases in fission yeast by distinct cellular localization and in vivo specificity.

Funktionelle Unterschiede zwischen Histondeacetylasen in Spalthefe durch unterschiedliche zellul&#xE4;re Lokalisation und In-vivo-Spezifit&#xE4;t.

A transcriptionally silent chromosomal domain is found in the mating-type region of fission yeast. Here we show that this domain is delimited by 2-kb inverted repeats, IR-L and IR-R. IR-L and IR-R prevent the expansion of transcription-permissive chromatin into the silenced region and that of silenced chromatin into the expressed region. Their insulator activity is partially orientation dependent. The silencing defects that follow deletion or inversion of IR-R are suppressed by high dosage of the chromodomain protein Swi6. Combining chromosomal deletions and Swi6 overexpression shows that IR-L and IR-R provide firm borders in a region where competition between silencing and transcriptional competence occurs. IR-R possesses autonomously replicating sequence (ARS) activity, leading to a model where replication factors, or replication itself, participate in boundary formation.

Expression-state boundaries in the mating-type region of fission yeast.

Expression-staatlichen Grenzen in der Paarungszeit-Typ-Region von S. pombe.

Hrp3, a paralog of Hrp1, is a novel member of the CHD1 (chromo-helicase/ATPase-DNA binding 1) protein family of Schizosaccharomyces pombe. Although it has been considered that CHD1 proteins are required for chromatin modifications in transcriptional regulations, little is known about their roles in vivo. In this study, we examined the effects of Hrp3 on heterochromatin silencing using several S. pombe reporter strains. The phenotypic analysis revealed that hrp3(+) is not an essential gene for cell viability. However, Hrp3 is required for transcriptional repression at silence loci of mat3. A chromatin immunoprecipitation assay showed that Hrp3 directly associates with mat3 chromatin. Thus, our results strongly suggest that Hrp3 is involved in heterochromatin silencing and plays a direct role as a chromatin remodeling factor at mat3 in vivo.

Hrp3, a chromodomain helicase/ATPase DNA binding protein, is required for heterochromatin silencing in fission yeast.

Hrp3, ein Chromodom&#xE4;ne Helikase / ATPase DNA-bindendes Protein, ist f&#xFC;r Heterochromatin Silencing in Spalthefe erforderlich.

The post-translational modifications of histones are key to the modulation of chromatin structure. Distinct patterns of modifications established by histone-modifying enzymes control diverse chromosomal processes. Here, we report the purification and molecular characterization of the fission yeast Clr6 histone deacetyl ase involved in higher order chromatin assembly. We show that a chromodomain protein Alp13, which belongs to the conserved MRG protein family linked to cellular senescence in humans, is associated with Clr6. In addition, Clr6 interacts with homologs of the mammalian transcriptional co-repressors Sin3, Pst1 and Pst2, and a WD40 repeat-containing protein, Prw1. Alp13, Pst2 and Prw1 form a stable complex with Clr6 in the nucleus. Deletion of any of these factors causes progressive loss of viability and sensitivity to DNA-damaging agents, and impairs condensation/resolution of chromosomes during mitosis. This is accompanied by hyperacetylation of histones and a reduction in histone H3 Ser10 phosphorylation, which correlates with chromosome condensation during mitosis. These results link the MRG family protein Alp13 to histone deacetylation, and suggest that Clr6 and its associated factors are essential for fundamental chromosomal events.

Alp13, an MRG family protein, is a component of fission yeast Clr6 histone deacetylase required for genomic integrity.

Alp13, ein Protein MRG-Familie, ist eine Komponente des Spalthefe Clr6 Histondeacetylase f&#xFC;r genomischen Integrit&#xE4;t erforderlich.

The mating-type region of the fission yeast Schizosaccharomyces pombe comprises three loci: mat1, mat2-P and mat3-M. mat1 is expressed and determines the mating type of the cell. mat2-P and mat3-M are two storage cassettes located in a 17 kb heterochromatic region with features identical to those of mammalian heterochromatin. Mutations in the swi6+, clr1+, clr2+, clr3+, clr4+ and clr6+ genes were obtained in screens for factors necessary for silencing the mat2-P-mat3-M region. swi6+ encodes a chromodomain protein, clr3+ and clr6+ histone deacetylases, and clr4+ a histone methyltransferase. Here, we describe the cloning and characterization of clr2+. The clr2+ gene encodes a 62 kDa protein with no obvious sequence homologs. Deletion of clr2+ not only affects transcriptional repression in the mating-type region, but also centromeric silencing and silencing of a PolII-transcribed gene inserted in the rDNA repeats. Using chromatin immunoprecipitation, we show that Clr2 is necessary for histone hypoacetylation in the mating-type region, suggesting that Clr2 acts upstream of histone deacetylases to promote transcriptional silencing.

A novel type of silencing factor, Clr2, is necessary for transcriptional silencing at various chromosomal locations in the fission yeast Schizosaccharomyces pombe.

Ein neuartiger Typ von Silencing-Faktor, Clr2, ist notwendig f&#xFC;r transkriptionellen Silencing an verschiedenen Orten in der chromosomalen Spalthefe Schizosaccharomyces pombe.

Centromeres of fission yeast are arranged with a central core DNA sequence flanked by repeated sequences. The centromere-associated histone H3 variant Cnp1 (SpCENP-A) binds exclusively to central core DNA, while the heterochromatin proteins and cohesins bind the surrounding outer repeats. CHD (chromo-helicase/ATPase DNA binding) chromatin remodeling factors were recently shown to affect chromatin assembly in vitro. Here, we report that the CHD protein Hrp1 plays a key role at fission yeast centromeres. The hrp1Delta mutant disrupts silencing of the outer repeats and central core regions of the centromere and displays chromosome segregation defects characteristic for dysfunction of both regions. Importantly, Hrp1 is required to maintain high levels of Cnp1 and low levels of histone H3 and H4 acetylation at the central core region. Hrp1 interacts directly with the centromere in early S-phase when centromeres are replicated, suggesting that Hrp1 plays a direct role in chromatin assembly during DNA replication.

The CHD remodeling factor Hrp1 stimulates CENP-A loading to centromeres.

Die CHD Umbau Faktor HRP1 stimuliert CENP-A Laden zu Centromeren.

The genome has a non-random spatial distribution in the cell nucleus. In Schizosaccharomyces pombe, it has been shown that the centromeres, telomeres and the mating-type region localize to the nuclear membrane (NM), the former by attaching to the spindle pole body (SPB). In addition, reporter genes inserted into these areas are transcriptionally repressed because of the formation of specialized chromatin structures. Performing live cell analysis we found that in a wild-type strain the mating-type region was positioned in the proximity of the SPB, the location where the pericentromeric heterochromatin is also found. In a strain lacking the histone methyltransferase Clr4, crucial for the formation of heterochromatin, the mating-type region had a random localization in the nucleus. Moreover, in a strain in which the two boundary elements IR-L and IR-R had been deleted, the mating-type region was displaced from its position at the proximity of the SPB, but remained in the vicinity of the NM. Moreover, in all investigated strains with silencing deficiencies the distance between the mating-type region and the SPB increased. This result indicates a correlation between transcriptional derepression and displacement of the region. Two different models of how the mating-type chromatin is organized in the nucleus are discussed.

The Clr4 methyltransferase determines the subnuclear localization of the mating-type region in fission yeast.

Das Genom hat eine nicht zuf&auml;llige r&auml;umliche Verteilung im Zellkern. In Schizosaccharomyces Pombe hat sich gezeigt, dass die Zentromerzahl, Telomere und die Paarung-Typ-Region um die Kernmembrane (NM), erstere durch Anf&uuml;gen der Spindel-Pol-Stelle (SRA) zu lokalisieren. Dar&uuml;ber hinaus werden Reporter Gene in diesen Bereichen eingef&uuml;gt transcriptionally wegen der Bildung von spezialisierten Chromatin Strukturen unterdr&uuml;ckt. Durchf&uuml;hren von live Zellanalyse fanden wir, dass in einem Wildtyp-Stamm die Paarung-Typ-Region in der N&auml;he von SPB, die Lage positioniert war, die perizentromerische Heterochromatin wird auch festgestellt. Eine Sorte fehlen die Histon-Methyltransferase Clr4, entscheidend f&uuml;r die Bildung von Heterochromatin hatte die Paarung-Typ-Region eine zuf&auml;llige Lokalisierung im Zellkern. Dar&uuml;ber hinaus wurde eine Belastung, in der die zwei Grenze Elemente IR-L und IR-R gel&ouml;scht worden, die Paarung-Typ-Region aus seiner Position in der N&auml;he von der SPB vertrieben, blieb aber in der N&auml;he der NM. Dar&uuml;ber hinaus untersuchten in allen St&auml;mme mit Unterdr&uuml;ckender M&auml;ngel, die der Abstand zwischen dem Paarung-Typ und das SPB erh&ouml;ht. Dieses Ergebnis deutet auf eine Korrelation zwischen transkriptionelle Derepression und Verschiebung der Region. Zwei verschiedene Modelle der Anordnung von des Paarung-Typ-Chromatins in den Zellkern werden diskutiert.

Die Clr4-Methyltransferase bestimmt die Elementarpartikel Lokalisierung der Region Paarung-Typ im Bereich der Kernspaltung Hefe.

There are several documented events of changes in subnuclear localization during gene activation. However, there are conflicting data on whether the nuclear periphery is a compartment for gene repression or activation and whether genes are moved to the pores at the nuclear membrane (NM) or not during gene activation. Nitrogen starvation of fission yeast serves as a good model system for studying gene induction, as it causes fast regulation of hundreds of genes. In this study, the subnuclear localization of two gene clusters repressed by nitrogen was investigated. During normal growth conditions, the gene clusters localized to the nuclear periphery at the opposite side of the nucleus as compared to the spindle pole body. This constrained localization was dependent on the histone deacetylase Clr3, known to transcriptionally repress genes in these clusters. Already 20 min after nitrogen depletion, drastic changes in subnuclear localization of the two loci were observed, away from the NM toward the nuclear interior. At least for one of the clusters, the movement was clearly transcription dependent. Data presented in this paper illustrates how interconnected events of gene activation and nuclear reorganization are as well as provides a suggestion of how nuclear organization might be maintained.

Reorganization of chromatin is an early response to nitrogen starvation in Schizosaccharomyces pombe.

Es gibt mehrere dokumentierte Ereignisse von &Auml;nderungen in der Elementarpartikel Lokalisierung w&auml;hrend der Genaktivierung. Allerdings gibt es widerspr&uuml;chliche Daten, ob die nukleare Peripherie ist ein Fach f&uuml;r gen Repression oder Aktivierung und ob Gene in die Poren auf der Kernmembrane (NM) oder verschoben werden nicht w&auml;hrend der Genaktivierung. Stickstoff Hungertod von Kernspaltung Hefe dient als gutes Modellsystem zur Untersuchung von Gen-Induktion, wie es schnelle Regulierung der Hunderte von Genen verursacht. In dieser Studie wurde untersucht, die Elementarpartikel Lokalisierung von zwei Gen-Cluster unterdr&uuml;ckt durch Stickstoff. W&auml;hrend der normalen Wachstumsbedingungen lokalisiert die Gen-Cluster zur nuklearen Peripherie an der gegen&uuml;berliegenden Seite des Kerns im Vergleich zu der Spindel-Pol-K&ouml;rper. Diese eingeschr&auml;nkte Lokalisierung war abh&auml;ngig von der Histon-Deacetylase Clr3, bekannt, dass Gene in diesen Haufen transcriptionally zu unterdr&uuml;cken. Bereits 20 min nach Stickstoff Ersch&ouml;pfung, Richtungs&auml;nderungen Elementarpartikel Lokalisierung der zwei Loci, Weg von der NM gegen&uuml;ber der nuklearen Innenraum beobachtet. Zumindest war die Bewegung einerseits der Cluster eindeutig Transkription angewiesen. Daten pr&auml;sentiert in diesem Papier werden wie miteinander verbundene Ereignisse der Genaktivierung und nukleare Reorganisation sind als auch erm&ouml;glicht, dass ein Vorschlag wie nukleare Organisation aufrechterhalten werden k&ouml;nnte.

Reorganisation des Chromatins ist eine fr&uuml;he Antwort auf Stickstoff verhungern in Schizosaccharomyces Pombe.

Tripeptidyl-peptidase II (TPP II) is a cytosolic peptidase that has been implicated in fat formation and cancer, apparently independent of the enzymatic activity. In search for alternative functional regions, conserved motifs were identified and eleven signatures were constructed. Seven of the signatures covered previously investigated residues, whereas the functional importance of the other motifs is unknown. This provides directions for future investigations of alternative activities of TPP II. The obtained signatures provide an efficient bioinformatic tool for the identification of TPP II homologues. Hence, a TPP II sequence homologue from fission yeast, Schizosaccharomyces pombe, was identified and demonstrated to encode the TPP II-like protein previously reported as multicorn. Furthermore, an homologous protein was found in the prokaryote Blastopirellula marina, albeit the TPP II function was apparently not conserved. This gene is probably the result of a rare gene transfer from eukaryote to prokaryote.

Development, evaluation and application of tripeptidyl-peptidase II sequence signatures.

Tripeptidyl Peptidase II (TPP II) ist ein Cytosolisches Peptidase, die in Fett Bildung und Krebs, offenbar unabh&auml;ngig von der Enzymaktivit&auml;t verwickelt hat. Auf der Suche nach alternativen funktionale Regionen konservierte Motive wurden identifiziert und elf Unterschriften wurden gebaut. Sieben der Signaturen zuvor erfassten untersucht R&uuml;ckst&auml;nde, w&auml;hrend die funktionelle Bedeutung der anderen Motive unbekannt ist. Dies enth&auml;lt Anweisungen f&uuml;r zuk&uuml;nftige Untersuchungen zu alternativen T&auml;tigkeiten TPP-II. Die erhaltenen Signaturen bieten ein effizientes Bioinformatic Hilfsmittel zur Identifizierung von TPP II Homologen. Daher wurde ein TPP II-Sequenz-Homolog von Kernspaltung Hefe Schizosaccharomyces Pombe, identifiziert und demonstriert das TPP II-wie Protein, die zuvor als Multicorn gemeldet zu codieren. Au&szlig;erdem wurde eine homologe Protein in der Prokaryoten Blastopirellula Marina, festgestellt, wenn auch die TPP II-Funktion anscheinend nicht konserviert wurde. Dieses Gen ist wahrscheinlich das Ergebnis einer seltenen Gentransfers von Eukaryoten auf Prokaryoten.

Erarbeitung, Bewertung und Anwendung von Tripeptidyl Peptidase II Sequenz Unterschriften.

Gene transcription is associated with local changes in chromatin, both in nucleosome positions and in chemical modifications of the histones. Chromatin dynamics has mostly been studied on a single-gene basis. Those genome-wide studies that have been made primarily investigated steady-state transcription. However, three studies of genome-wide changes in chromatin during the transcriptional response to heat shock in the budding yeast Saccharomyces cerevisiae revealed nucleosome eviction in promoter regions but only minor effects in coding regions. Here, we describe the short-term response to nitrogen starvation in the fission yeast Schizosaccharomyces pombe. Nitrogen depletion leads to a fast induction of a large number of genes in S. pombe and is thus suitable for genome-wide studies of chromatin dynamics during gene regulation. After 20 min of nitrogen removal, 118 transcripts were up-regulated. The distribution of regulated genes throughout the genome was not random; many up-regulated genes were found in clusters, while large parts of the genome were devoid of up-regulated genes. Surprisingly, this up-regulation was associated with nucleosome eviction of equal magnitudes in the promoters and in the coding regions. The nucleosome loss was not limited to induction by nitrogen depletion but also occurred during cadmium treatment. Furthermore, the lower nucleosome density persisted for at least 60 min after induction. Two highly induced genes, urg1(+) and urg2(+), displayed a substantial nucleosome loss, with only 20% of the nucleosomes being left in the coding region. We conclude that nucleosome loss during transcriptional activation is not necessarily limited to promoter regions.

Nitrogen depletion in the fission yeast Schizosaccharomyces pombe causes nucleosome loss in both promoters and coding regions of activated genes.

Gen-Transkription ist mit lokalen &Auml;nderungen in Chromatin, beide in Nukleosom Positionen und chemische Modifikationen des die Histone verbunden. Chromatin Dynamik wurde &uuml;berwiegend auf Basis von Einzel-gen untersucht. Diese genomweiten-Studien, die in erster Linie vorgenommen wurden untersucht station&auml;ren Transkription. Drei Studien der genomweiten &Auml;nderungen in Chromatin w&auml;hrend der transcriptional Antwort auf Hitzeschock in der Backhefe ergab jedoch Nukleosom R&auml;umung in Projekttr&auml;ger Regionen aber nur geringf&uuml;gige Auswirkungen in proteinkodierenden Regionen. Hier beschreiben wir die kurzfristige Reaktion auf Stickstoff verhungern in der Kernspaltung Hefe Schizosaccharomyces Pombe. Stickstoff Ersch&ouml;pfung f&uuml;hrt zu eine schnelle Induktion einer gro&szlig;en Anzahl von Genen in S. Pombe und eignet sich somit f&uuml;r genomweiten Studien der Chromatin-Dynamik w&auml;hrend der Genregulation. Nach 20 min von Stickstoff-Entfernung wurden 118 Transkripte oben-geregelt. Die Verteilung der regulierte Gene in das Genom wurde nicht zuf&auml;llig; viele oben-geregelt-Gene wurden in Clustern, gefunden, w&auml;hrend gro&szlig;e Teile des Genoms frei von Genen oben-geregelt waren. &Uuml;berraschenderweise wurde dieser bis-Verordnung Nukleosom R&auml;umung gleicher Gr&ouml;&szlig;enordnung in den Projekttr&auml;gern und proteinkodierenden Regionen zugeordnet. Der Nukleosom-Verlust war nicht beschr&auml;nkt auf Induktion von Stickstoff Ersch&ouml;pfung aber auch Kadmium Behandlung aufgetreten. Dar&uuml;ber hinaus beibehalten die geringere Dichte Nukleosom f&uuml;r mindestens 60 Minuten nach Induktion. Zwei hoch induzierte Genen, urg1(+) und urg2(+), angezeigt einen erheblichen Nukleosom-Verlust, mit nur 20 % der Nucleosomes in der Kodierungregion &uuml;berlassen. Wir schlie&szlig;en, dass Nukleosom Verlust w&auml;hrend der transcriptional Aktivierung nicht unbedingt Projekttr&auml;ger Regionen beschr&auml;nkt.

Stickstoff-Abbau in der Kernspaltung Hefe Schizosaccharomyces Pombe verursacht Nukleosom Verlust Promotoren sowohl kodierende Regionen der aktivierten Gene.

Significant progress has been made in understanding the functional organisation of the cell nucleus. Still many questions remain to be answered about the relationship between the spatial organisation of the nucleus and the regulation of the genome function. There are many conflicting data in the field making it very difficult to merge published results on mammalian cells into one model on subnuclear chromatin organisation. The fission yeast, Schizosaccharomyces pombe, over the last decades has emerged as a valuable model organism in understanding basic biological mechanisms, especially the cell cycle and chromosome biology. In this review we describe and compare the nuclear organisation in mammalian and fission yeast cells. We believe that fission yeast is a good tool to resolve at least some of the contradictions and unanswered questions concerning functional nuclear architecture, since S. pombe has chromosomes structurally similar to that of human. S. pombe also has the advantage over higher eukaryotes in that the genome can easily be manipulated via homologous recombination making it possible to integrate the tools needed for visualisation of chromosomes using live-cell microscopy. Classical genetic experiments can be used to elucidate what factors are involved in a certain mechanism. The knowledge we have gained during the last few years indicates similarities between the genome organisation in fission yeast and mammalian cells. We therefore propose the use of fission yeast for further advancement of our understanding of functional nuclear organisation.

Advancing our understanding of functional genome organisation through studies in the fission yeast.

Zum Verst&auml;ndnis der funktionellen Organisation des Zellkerns wurden erhebliche Fortschritte erzielt. Noch viele Fragen bleiben um &uuml;ber die Beziehung zwischen der r&auml;umlichen Organisation des Zellkerns und der Regelung der Genom-Funktion beantwortet werden. Es gibt viele widerspr&uuml;chliche Daten im Feld, so dass es sehr schwierig, ein Modell auf Elementarpartikel Chromatin Organisation ver&ouml;ffentlichten Ergebnisse auf S&auml;ugetier-Zellen zusammenf&uuml;hren. Die Kernspaltung-Hefe Schizosaccharomyces Pombe, als eine wertvolle Modellorganismus in der grundlegenden biologischen Mechanismen Verst&auml;ndnis, vor allem den Zellzyklus und Chromosom Biologie in den letzten Jahrzehnten entstanden ist. In diesem Bericht wir beschreiben und vergleichen die nukleare Organisation in S&auml;ugetier- und Kernspaltung Hefezellen. Wir glauben, dass die Kernspaltung Hefe ist ein gutes Werkzeug, um zumindest einige Aufl&ouml;sen der Widerspr&uuml;che und unbeantwortete Fragen zur funktionalen nukleare Architektur, seit S. Pombe hat strukturell &auml;hnlich der menschlichen Chromosomen. S. Pombe hat auch den Vorteil gegen&uuml;ber h&ouml;heren Eukaryotes, da das Genom durch homologe Rekombination macht es m&ouml;glich, die zur Visualisierung der Chromosomen mit Leben-Zelle Mikroskopie erforderlichen Tools integrieren leicht manipuliert werden kann. Klassische genetische Experimente k&ouml;nnen verwendet werden, um herauszustellen, welche Faktoren in einem bestimmten Mechanismus beteiligt sind. Das wissen, das wir in den letzten Jahren gewonnen haben zeigt &Auml;hnlichkeiten zwischen der Genom-Organisation in der Kernspaltung Hefe und S&auml;ugetier-Zellen. Wir schlagen deshalb vor, die Nutzung der Kernspaltung Hefe f&uuml;r das weitere Fortkommen unseres Verst&auml;ndnisses von funktionellen nukleare Organisation.

Article	Year
Functional divergence between histone deacetylases in fission yeast by distinct cellular localization and in vivo specificity. Molecular and cellular biology\| PubMed ID: 11884604	2002
Expression-state boundaries in the mating-type region of fission yeast. Genetics\| PubMed ID: 12072458	2002
Hrp3, a chromodomain helicase/ATPase DNA binding protein, is required for heterochromatin silencing in fission yeast. Biochemical and biophysical research communications\| PubMed ID: 12127990	2002
Alp13, an MRG family protein, is a component of fission yeast Clr6 histone deacetylase required for genomic integrity. The EMBO journal\| PubMed ID: 12773392	2003
A novel type of silencing factor, Clr2, is necessary for transcriptional silencing at various chromosomal locations in the fission yeast Schizosaccharomyces pombe. Nucleic acids research\| PubMed ID: 15317867	2004
The CHD remodeling factor Hrp1 stimulates CENP-A loading to centromeres. Nucleic acids research\| PubMed ID: 15908586	2005
The Clr4 methyltransferase determines the subnuclear localization of the mating-type region in fission yeast. Journal of cell science\| PubMed ID: 17504808	2007
Reorganization of chromatin is an early response to nitrogen starvation in Schizosaccharomyces pombe. Chromosoma\| PubMed ID: 18936951	2009
Development, evaluation and application of tripeptidyl-peptidase II sequence signatures. Archives of biochemistry and biophysics\| PubMed ID: 19467630	2009
Nitrogen depletion in the fission yeast Schizosaccharomyces pombe causes nucleosome loss in both promoters and coding regions of activated genes. Genome research\| PubMed ID: 20086243	2010
Advancing our understanding of functional genome organisation through studies in the fission yeast. Current genetics\| PubMed ID: 21113595	2011

Pernilla Bjerling

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