Phylogenetic analyses of penicillia based on partial calmodulin gene sequences

Partial sequences (about 600 nucleotides) of the calmodulin gene were used for the phylogenetic studies on Eupenicillium, Talaromyces and Penicillium. This region is from the 3rd base of the codon for the 9th amino acid Gln to the 3rd base of the codon for the 122th amino acid Val, flanking parts of the 2nd and 5th exons with complete sequences of two exons and three introns. Seventy-six isolates of 56 taxa of penicillia were involved. The nucleotide sequences with and without introns were analyzed respectively using the neighbor-joining (NJ) and maximum parsimony (MP) methods. The cluster analysis on relative synonymous codon usage (RSCU) of each sequence was also carried out. The fact that species of penicillia belong to the two subfamilies of the Trichocomaceae proposed by Malloch based on traditional methods is supported by our molecular data, whereas, the development of asci and patterns of penicilli show little phylogenetic information. Nine groups in the lineage of Eupenicillium and two in that of Talaromyces were recognized in our studies. In addition to the teleomorph-holomorph-anamorph evolutionary model of penicillia suggested by LoBuglio et al., and Pitt, we proposed that a mutation bias of holomorphs/anamorphs with or without selection is another evolutionary path of these organisms.

[1]  D. T. Wicklow,et al.  Ribosomal RNA Comparisons among Taxa of the Terverticillate Penicillia , 1990 .

[2]  Xiao Sun,et al.  Cluster analysis of the codon use frequency of MHC genes from different species. , 2002, Bio Systems.

[3]  B. O. Dodge The Perithecium and Ascus of Penicillium , 1933 .

[4]  Jens Christian Frisvad,et al.  Phylogenetic analysis of Penicillium subgenus Penicillium using partial P-tubulin sequences , 2004 .

[5]  Charles Thom,et al.  A Manual of the Penicillia , 1949 .

[6]  D. Lynn,et al.  Synonymous codon usage is subject to selection in thermophilic bacteria. , 2002, Nucleic acids research.

[7]  R. Samson,et al.  Studies on Talaromyces and related genera I. Hamigera gen. nov. and Byssochlamys , 1971 .

[8]  David Malloch,et al.  The Trichocomaceae: Relationships with Other Ascomycetes , 1986 .

[9]  D. Reynolds,et al.  The fungal holomorph : mitotic, meiotic and pleomorphic speciation in fungal systematics : proceedings of an International Symposium, Newport, Oregon, 4-7 August, 1992 , 1993 .

[10]  Francisco Infante,et al.  Penicillium brocae, a new species associated with the coffee berry borer in Chiapas, Mexico. , 2003, Mycologia.

[11]  J. Frisvad,et al.  Chemotaxonomy of the genus Talaromyces , 1990, Antonie van Leeuwenhoek.

[12]  Chester R. Benjamin,et al.  Ascocarps of Aspergillus and Penicillium , 1955 .

[13]  J. Pitt The genus Penicillium and its teleomorphic states Eupenicillium and Talaromyces , 1981 .

[14]  John I. Pitt,et al.  Modern Concepts in Penicillium and Aspergillus Classification , 1992, NATO ASI Series.

[15]  Dorothy I. Fennell,et al.  The Genus Eupenicillium Ludwig , 1969 .

[16]  Jens Christian Frisvad,et al.  List of accepted species and their synonyms in the family Trichocomaceae , 2000 .

[17]  J. I. Pitt,et al.  Phylogeny in the genus Penicillium: a morphologist's perspective , 1995 .

[18]  P. Sharp,et al.  Synonymous codon usage in Saccharomyces cerevisiae , 1991, Yeast.

[19]  Robert A. Samson,et al.  The ascomycete genus eupenicillium and related penicillium anamorphs , 1983 .

[20]  Lynne Sigler,et al.  Four new Penicillium species having Thysanophora -like melanized conidiophores , 2002 .

[21]  A. C. Stolk,et al.  Studies on the genusEupenicillium ludwig III. Four new species ofEupenicillium , 2005, Antonie van Leeuwenhoek.

[22]  B. Hall,et al.  Phylogenetic relationships among ascomycetes: evidence from an RNA polymerse II subunit. , 1999, Molecular biology and evolution.

[23]  S. Carroll,et al.  Genome-scale approaches to resolving incongruence in molecular phylogenies , 2003, Nature.

[24]  Robert A. Samson,et al.  Penicillium persicinum, a new griseofulvin, chrysogine and roquefortine C producing species from Qinghai province, China , 2004, Antonie van Leeuwenhoek.

[25]  Paul M. Sharp,et al.  Codon usage in yeast: cluster analysis clearly differentiates highly and lowly expressed genes , 1986, Nucleic Acids Res..

[26]  Peden Jf,et al.  Analysis of codon usage. , 2000 .

[27]  Jens Christian Frisvad,et al.  Fungal identification techniques , 1996 .

[28]  T. A. Hall,et al.  BIOEDIT: A USER-FRIENDLY BIOLOGICAL SEQUENCE ALIGNMENT EDITOR AND ANALYSIS PROGRAM FOR WINDOWS 95/98/ NT , 1999 .

[29]  J. Thompson,et al.  The CLUSTAL_X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. , 1997, Nucleic acids research.

[30]  J. Felsenstein Cases in which Parsimony or Compatibility Methods will be Positively Misleading , 1978 .

[31]  Long Wang,et al.  DESIGNING PRIMER SETS FOR AMPLIFICATION OF PARTIAL CALMODULIN GENES FROM PENICILLIA , 2004 .

[32]  N. L. Glass,et al.  Development of primer sets designed for use with the PCR to amplify conserved genes from filamentous ascomycetes , 1995, Applied and environmental microbiology.

[33]  M. Berbee,et al.  Fungal Molecular Evolution: Gene Trees and Geologic Time , 2001 .

[34]  John I. Pitt,et al.  Integration of modern taxonomic methods for Penicillium and Aspergillus classification , 2000 .

[35]  M. Gouy,et al.  Codon catalog usage and the genome hypothesis. , 1980, Nucleic acids research.

[36]  D. B. Scott,et al.  Studies on the genusEupenicillium Ludwig II. Perfect states of some Penicillia , 2005, Antonie van Leeuwenhoek.

[37]  C. W. Emmons The Ascocarps in Species of Penicillium , 1935 .

[38]  John I. Pitt,et al.  Advances in Penicillium and Aspergillus Systematics , 1986, NATO ASI Series.

[39]  S. Peterson,et al.  Multilocus DNA sequence analysis shows that Penicillium biourgeianum is a distinct species closely related to P. brevicompactum and P. olsonii. , 2004, Mycological research.

[40]  Ning Zhang,et al.  Molecular phylogeny of Melanospora and similar pyrenomycetous fungi , 2002 .

[41]  J. Sugiyama,et al.  Is Penicillium monophyletic? An evaluation of phylogeny in the family Trichocomaceae from 18S, 5.8S and ITS ribosomal DNA sequence data. , 1995 .

[42]  Ignazio Carbone,et al.  A method for designing primer sets for speciation studies in filamentous ascomycetes , 1999 .

[43]  J. Sugiyama,et al.  Polyphyletic origins of species of the anamorphic genus Geosmithia and the relationships of the cleistothecial genera: Evidence from 18S, 5S and 28S rDNA sequence analyses , 1997 .

[44]  J. Frisvad,et al.  Reclassification of the Penicillium roqueforti group into three species on the basis of molecular genetic and biochemical profiles. , 1996, Microbiology.

[45]  J. Pitt,et al.  Phylogenetic analysis of two ribosomal DNA regions indicates multiple independent losses of a sexual , 1993 .

[46]  Jens Christian Frisvad,et al.  Introduction to food- and airborne fungi. , 2004 .

[47]  Jens Christian Frisvad,et al.  Systematics of Penicillium simplicissimum based on rDNA sequences, morphology and secondary metabolites , 2001 .

[48]  Jens Christian Frisvad,et al.  Penicillium subgenus Penicillium - A guide to identification of food and air-borne terverticillate Penicillia and their mycotoxins , 2004 .

[49]  John I. Pitt,et al.  Independent origins of the synnematous Penicillium species, P. duclauxii, P. clavigerum, and P. vulpinum, as assessed by two ribosomal DNA regions , 1994 .

[50]  Ailsa D. Hocking,et al.  The Closely Related Species of Subgenus Penicillium — A Phylogenic Exploration , 1986 .

[51]  Jens Christian Frisvad,et al.  Molecular methods for differentiation of closely related Penicillium species , 2000 .

[52]  J. W. Taylor,et al.  Phylogeny and PCR identification of the human pathogenic fungus Penicillium marneffei , 1995, Journal of clinical microbiology.

[53]  D. B. Scott,et al.  Studies on the genus Eupenicillium Ludwig I. Taxonomy and nomenclature of Penicillia in relation to their sclerotioid ascocarpic states , 1967 .

[54]  Jens Christian Frisvad,et al.  Phylogenetic analysis of nucleotide sequences from the ITS region of terverticillate Penicillium species , 1999 .