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Comparative Genomics and Evolutionary Analysis of Cytochrome P450
Monooxygenases in Fungal Subphylum Saccharomycotina
Article  in  Journal of Pure and Applied Microbiology · November 2014
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Ipeleng Kgosiemang
Central University of Technology
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Comparative Genomics and Evolutionary Analysis of
Cytochrome P450 Monooxygenases in Fungal Subphylum
Saccharomycotina
Ipeleng Kopano Rosinah Kgosiemang,
Samson Sitheni Mashele and Khajamohiddin Syed*
Department of Health Sciences, Faculty of Health and Environmental Sciences,
Central University of Technology, Bloemfontein 9300, Free State, South Africa.
(Received: 09 September 2014; accepted: 29 October 2014)
Cytochrome P450 monooxygenases (P450s) are heme-thiolate enzymes and play
an important role in the primary and secondary metabolism of living organisms. Genome
sequencing analysis of fungal organisms revealed the presence of numerous P450s in
their genomes, with few exceptions. P450s in the fungal subphylum Saccharomycotina,
which contains biotechnologically important and opportunistic human pathogen yeasts,
have been underexplored because there are few P450s in their genomes. In the present
study we performed comparative analysis of P450s in 25 yeast species. A hundred and
seventy-two P450s were found in 25 yeast species and these are grouped into 13 P450
families and 27 subfamilies. P450s ranged from a minimum of three (Saccharomyces
species) to a maximum of 21 (Candida species) in the yeast genomes. Among the P450
families, the CYP52 family showed the highest number of member P450s (71) followed by
CYP51 (27), CYP61 (25), CYP56 (20) and CYP501 (11). Pichia pastoris and Dekkera
bruxellensis showed a novel P450 family, CYP5489, in their genome.  Based on the
functional properties of characterized P450s, we conclude that P450s in Saccharomycotina
species possibly play a role in organisms’ physiology either in the synthesis of cellular
components or in the utilization of simpler organic molecules. The ecological niches of
yeast species are highly enriched with simpler organic nutrients and it is well known
that yeast species utilize simpler organic nutrients as carbon source efficiently. This
might have played a role in compacting yeast genomes and possibly losing a considerable
number of P450s during evolution.
Key words: Adaptation, Comparative genomics, Cytochrome P450 monooxygenases,
 Ecological niches, Evolution, Fungi, Saccharomycotina, Yeast.
The fungal kingdom is the largest
biological kingdom and consists of diverse
organisms that are adapted to diverse ecological
niches. This kingdom is classified into four phyla,
namely Ascomycota, Basidiomycota, Zygomycota
and Chytridiomycota1. The phylum Ascomycota
is further classified into three subphyla, namely
Pezizomycotina, Saccharomycotina and
Taphrinomycotina2. Our interest lies in the
Saccharomycotina subphylum, which comprises
of  yeast organisms that are well known for their
potential biotechnological value, as well as
opportunistic human pathogens. Table 1 shows a
list of Saccharomycotina species, provides general
information and indicates the importance of these
species.
Genome sequencing analysis of fungal
species revealed the presence of numerous
cytochrome P450 monooxygenases (P450s) in their
genomes, with few exceptions3. P450s are heme-
thiolate enzymes and their role in organisms’
primary and secondary metabolism and their
potential use in biotechnology, bioremediation,
pharmacology and biofuel generation has been
J PURE APPL MICROBIO, 8(SPL. EDN.), NOVEMBER 2014.
292 KGOSIEMANG et al.:  P450OME OF SACCHAROMYCOTINA
documented4. Fungal P450s occupy a special place
because of their diverse catalytic activities
compared to the P450s from other biological
kingdoms5.  Furthermore, the presence of large
numbers of P450s and diverse families in fungi
makes fungal P450s ideal for the study of P450s’
evolution. Since the genome sequence of fungi
became available6, study on fungal P450s,
particularly on functional and evolutionary
aspects, has gained momentum. Study on fungal
P450s revealed that basidiomycetes enriched P450s
in their genome owing to duplication to help the
organism to adapt to diverse ecological niches,
such as wood degradation3,7. Study also revealed
that ascomycetes, particularly species belonging
to the subphylum Pezizomycotina, contain diverse
P450 families in their genome8. Furthermore, fungal
P450s were used as model P450s to identify P450
family-specific signature sequences9. Authors
have suggested that these signature sequences
evolved during the evolution of P450 families from
a common P450 ancestor. Recent studies also
revealed the presence of a large number of
thermostable P450s in fungi10. P450 evolutionary
studies or genome-wide comparative P450 studies
on fungal P450s were based on P450s from species
belonging to the phyla Basidiomycotoa and
Ascomycota. In the phylum Ascomycota only
P450s from the subphylum Pezizomycotina have
been explored for both evolutionary and functional
characterization3,8. P450s from Saccharomycotina
species are under explored and to date no study
on comparative P450s in Saccharomycotina
species has been carried out. Studies have been
limited to describing the number of P450s in
Saccharomycotina species and detailing one or two
P450 families. Furthermore, genome sequencing of
new Saccharomycotina species necessitated the
performance of comparative P450 profiling in their
genomes. Considering the importance of P450s in
general and the use of Saccharomycotina species
in biotechnology and their pathogenicity towards
animals, particularly humans, it is important to
understand the role of P450s in yeast species. In
the present study we performed genome-wide
comparative P450s analysis in 25 yeast species
and P450s in three yeast species, Pichia pastoris,
Dekkera bruxellensis and Pichia anomala, were
identified and annotated as per International
Cytochrome P450 Nomenclature Committee11.
MATERIALS   AND  METHODS
Saccharomycotina species
A total of 25 Saccharomycotina species
whose genomes have been sequenced and are
publicly available were used in the study. A list of
the yeast species used in this study is shown in
Table 1.
Genome-data mining of Saccharomycotina species
for P450s
The 22 Saccharomycotina species P450s
were downloaded from the publicly available
Cytochrome P450 Homepage12.  The downloaded
P450s were cross checked at the web pages: http:/
/www.yeastgenome.org/13 and http://
www.candidagenome.org/14. P450s in the remaining
three yeast species,i.e. P. pastoris, D. bruxellens
and P. anomala, were annotated following the
standard procedure described in our recent
studies7,10. Briefly, the proteome of yeast species
was downloaded from their genome data base
(Dekkera bruxellensis CBS 2499 v2.0: http://
genome.jgi.doe.gov/ Dekbr2/Dekbr2.home.html;
Pichia Pastoris: http://bioinformatics.
psb.ugent.be/orcae/;Pichia anomala[currently
named Wickerhamomyces anomalus]NRRL Y-366-
8 v1.0: http://genome.jgi.doe. gov/Wican1/
Wican1.home.html) and the whole proteome was
subjected to functional annotation using NCBI
Batch Web CD-search tool15. The proteins grouped
under the P450 superfamily were selected and
analyzed for the presence of the P450 family
signature motifs, namely EXXR and CXG. The
proteins that showed both motifs were considered
authentic P450s and used in this study.
Annotation and classification of P450s
The proteins identified as P450 were
subjected to BLAST analysis against all named
fungal species at the Cytochrome P450
Homepage12. For each P450, the closest homolog
was identified and based on the homology
percentage, family and subfamily names were
assigned. For assigning the family and subfamily
names, the standard rule set by the International
P450 Nomenclature Committee was followed, i.e.
P450s within a family share more than 40% amino
acid homology and members of subfamilies share
more than 55% amino acid homology11.
Furthermore, P450s that showed less than 40%
homology with known P450s were assigned to a
J PURE APPL MICROBIO, 8(SPL. EDN.), NOVEMBER 2014.
293KGOSIEMANG et al.:  P450OME OF SACCHAROMYCOTINA
new family with the help of the P450 nomenclature
of Dr David R Nelson, University of Tennessee
Health Science Center, Memphis, Tennessee, USA.
Phylogenetic analysis of P450s
Phylogenetic analysis of P450s was
carried out in the same way as described in our
recent publications7,10. Briefly, evolutionary
analysis was carried out using the minimum
evolution method16. The phylogenetic analysis
was carried out using Molecular Evolutionary
Genetics Analysis (MEGA 5.05) software17.
Functional analysis of P450s
Considering the large number of P450s
used in this study and the availability of functional
data, we performed a literature survey on the
functional analysis of Saccharomycotina species
P450s and their role in organisms’ physiology.
RESULTS   AND  DISCUSSION
P450ome of P. pastoris, D. bruxellens and P.
anomala
The yeast species P. pastoris and D.
bruxellens showed four P450s and P. anomala
showed six P450s in their genome (Table 2). Among
the P450 families found in these species CYP51,
CYP61 and CYP501 are common in three species.
Compared to two other yeast species, P450s
belonging to CYP56, CYP5205 and CYP5217
families are only present in P. anomala.
Surprisingly, P. pastoris and D. bruxellens showed
a novel P450 family, i.e. CYP5489, in their genome.
However, a difference was found in the CYP5489
subfamily type in yeast species.  P. pastoris
contained P450 belonging to subfamily A and D.
bruxellens contained P450 belonging to subfamily
B. It is noteworthy that this P450 family is not found
in other yeast species (Table 2).
Comparative analysis of P450ome in yeast species
A hundred and seventy-two P450s were
found in 25 yeast species (Table 2). The P450s
ranged from a minimum of three to a maximum of 21
in the yeast genomes; 172 P450s were grouped
under 13 P450 families and 27 subfamilies (Figure 1
and Table 2). Among the P450 families, CYP52
family showed the highest number of member P450s
(71) followed by CYP51 (27), CYP61 (25), CYP56
(20) and CYP501 (11) (Figure 2). The remaining P450
families showed less than 6% of member P450s
(Figure 2).
Fig. 1. Phylogenetic analysis of the P450s in subphyla Saccharomycotina. In total, 172 P450 sequences were
included in the tree. A minimal evolution tree was constructed using the close-neighbor-interchange algorithm in
MEGA (version 5.05). For ease of visual identity, P450s belonging to different families were presented in unique
colors.
J PURE APPL MICROBIO, 8(SPL. EDN.), NOVEMBER 2014.
294 KGOSIEMANG et al.:  P450OME OF SACCHAROMYCOTINA
Table 1.List of yeast species selected for the study and general information and importance of the species.
Species name General information and importance Reference
Saccharomyces The most useful yeast, which has been instrumental in winemaking, 24,25
cerevisiae  baking and brewing since ancient times. It is well known as baker’s
 yeast and it was the first eukaryote ever to have its genome
 completely sequenced. It is the microorganism behind the most
 common type of fermentation. It is one of the most intensively
 studied eukaryotic model organisms in molecular and cell biology,
 much like Escherichia coli as the model bacterium. It is the
 organism employed most in industry in terms of production volumes.
Saccharomyces Non-domesticated yeast living on the bark of deciduous trees. 26
paradoxus  Closest relative of S. cerevisiae hence it is an attractive model
 organism for the study of the population genetics and genomics of
 wild yeast.
Saccharomyces Belongs to Saccharomyces sensustricto complex. Shows high 26
mikatae  conservation of synteny with S. cerevisiae.
Saccharomyces Model organism for telomere research since many of the features of 27
castellii  the telomere biology of this yeast are closer to those of humans than
 other yeast.
Saccharomyces Used in winemaking, cider fermentation and making distilled 26, 27
bayanus  beverages. It is also used extensively for comparative genomic
 studies, including expression patterns and nucleosome profile.
Saccharomyces Belongs to Saccharomyces sensustricto complex. Used in making 27, 28
kudriavzevii  beer and wine. It can grow at low temperatures.
Saccharomyces More efficient user of glucose than model yeast S. cerevisiae. It is 27
kluyveri  used in industrial applications, such as the production of proteins, as
 its biomass yield is greater than that of S. cerevisiae. Lives in
 diverse environments, for example this yeast is a plant pathogen and
 has been isolated from drosophila species and tissues of an HIV-
infected patient.
Candida It is one of the major opportunistic pathogenic fungi causing 29, 30, 31
albicans  systemic infection (candidiasis) in immuno-compromised and
 immuno-competent hosts. It is a diploid fungus that grows both as
 yeast and filamentous cells and a casual agent of opportunistic oral
 and genital infections (superficial infections) in humans.
Candida The most prevalent opportunistic pathogenic yeast species of the 19, 31, 32
tropicalis Candida-non-albicans group. Human pathogen that is well known as
 medical yeast pathogen. It causes Candida-non-albican
 candidiasis. It is a glucose and maltose fermenting yeast. The
 industrial strain of C. tropicalis is engineered to produce long-chain-
dicarboxylic acids.
Candida The model organism of a group of so-called “flavinogenic yeasts” 32, 33
guilliermondii  capable of riboflavin oversynthesis. It often causes
(anamorph of  onychomycosis and is rarely involved in invasive fungal
Pichia guilliermondii)  infections.
Candida It was thought to be a primarily non-pathogenic organism of the 34
glabrata  normal flora of healthy individuals.   However, with the increasing
 population of immuno-compromised individuals, trends have shown
C. glabrata to be a highly opportunistic pathogen of the urogenital
 tract and of the bloodstream.
Candida A fungal species of the yeast family that has become a major cause 31
parapsilosis  of sepsis and of wound and tissue infections in immuno-
compromised patients. Among Candida species this yeast is a
 particular problem in neonates, transplant recipients and patients
J PURE APPL MICROBIO, 8(SPL. EDN.), NOVEMBER 2014.
295KGOSIEMANG et al.:  P450OME OF SACCHAROMYCOTINA
 receiving parenteral nutrition.
Candida It is the most closely related species to C. albicans and causes 35
dubliniensis  superficial infections. Compared to C. albicans, this yeast has low
 virulence and a longer survival rate in hosts.
Candida Responsible for less than 5% of invasive infections among Candida 31, 36
lusitaniae  species. Patients who undergo bone marrow transplantation and
 high-dose cytoreductive chemotherapy have been identified as being
 at risk of infections caused by this organism.
Kluyveromyces Best alternative yeast for genetics and physiology. Its name comes 37
lactis  from the ability to assimilate lactose and convert it to lactic acid.
 K. lactis is one of the main organisms grown in industry in
 fermenters to produce chymosin on a large scale. Chymosin is used
 for cheese production. This yeast has GRAS (generally regarded as
 safe) status.
Kluyveromyces Whole genome duplicated yeast. Distant lineage to S. cerevisiae and 38
polysporus  considered as best yeast to study comparative whole genome
 duplication event.
Kluyveromyces This yeast is a protoploid or pre whole genome duplication budding 39
waltii (recently  yeast descended from a lineage and did not undergo the whole
named  genome duplication event. Genome sequencing of this yeast
as  revealed how the genome of S. cerevisiae evolved via the whole
Lachancea waltii)  genome duplication event.
Pichia stipites Capable of both aerobic and oxygen-limited fermentation and has the 40
 highest known natural ability of any microbe to ferment xylose
 directly, converting it to ethanol.
Lodderomyces Causes blood stream infection in humans. It is the second or third 31
elongisporus  most common yeast species isolated from patients with bloodstream
 infections in Europe, Canada, Asia and Latin America.
Ashbya gossypii Contains the smallest genome of a free-living eukaryote and because 41
 of this is considered as model organism to study filamentous growth.
 It infects plants such as cotton and citrus. It is a natural overproducer
 of riboflavin (vitamin B2), which protects its spores against
 ultraviolet light. This makes it an interesting organism for industries,
 where genetically modified strains are still used to produce this
 vitamin.
Debaryomyces Metabolically versatile, osmotolerant and oleaginous. It is used for 31
hansenii  surface ripening of cheese and meat products. It one of the most
 halotolerant species capable of growing in media containing NaCl as
 high as 4 M.
Yarrowia Most extensively studied “non-conventional” yeast (strictly aerobic) 4, 43
lipolytica  and currently used as model organism for study of protein secretion,
peroxisome biogenesis, dimorphism, degradation of hydrophobic
substrates. It produces important metabolites and has developed
 intense secretory activity. Capable of growing on organic
 compounds and producing bio surfactants. This yeast has been used in
 industry (as a biocatalyst) and in academic studies. This yeast has
 GRAS  status.
Dekkera Although this yeast is distantly related to S. cerevisiae, is commonly 44
bruxellensis  found in the same habitat. It competes with S. cerevisiae in
 fermentation, can produce high amounts of ethanol and grow
 without oxygen. It plays a key role in flavor development in lambic
 beer. However, in the wine industry, this yeast is considered
 spoilage yeast owing to wine spoilage by the generation of a high
 amount of phenolic compounds (4-ethylguaiacol and 4-ethylphenol).
Table 1 Continue
J PURE APPL MICROBIO, 8(SPL. EDN.), NOVEMBER 2014.
296 KGOSIEMANG et al.:  P450OME OF SACCHAROMYCOTINA
Pichia This methylotrophic yeast is the most commonly used yeast species 45
pastoris  in the production of recombinant proteins and is widely used to
 produce proteins for basic research and medical applications. It is a
 model organism for the study of peroxisomal proliferation and
 methanol assimilation.
Wickerhamomyces It exhibits a multitude of biotechnologically important characteristics 46
anomalus  in flavor enhancement, food processing, biopreservation,
(formerly  dairy fermentation and waste water treatment.
Pichia anomala
and
Hansenula anomala)
Table 1 Continue
Among the 13 P450 families, CYP51 and
CYP61 were conserved across the 25 yeast species.
A single copy of CYP61 members were found in
the yeast species, whereas CYP51 members
were found in more than a single copy in
two yeast species , Candida guillermondii and
Kluyveromyces polysporus (Table 2).  The presence
of more than one copy of CYP51 members is not
rare and species belonging to other biological
kingdoms, such as plants, also showed more than
once copy of CYP51 members in their genomes18.
CYP52 family members are enriched in species
belonging to the genus Candida and in Yarrowia
lipolytica. Among candida species C. tropicalis
showed the maximum number of CYP52 members
(17 members) in its genome and Y. lipolytica
showed 12 members in its genome (Table 2).
Phylogenetic analysis of CYP52 members (Figure
1) resulted in the grouping of Y. lipolytica CYP52
members together, suggesting possible duplication
of member P450s in its genome after speciation.
Among yeast species only P. stipites,
Lodderomyces elongisporus and Debaryomyces
hansenii showed CYP52 members in their genome.
It is interesting to note that P. anomala did not
contain CYP52 members compared to P. stipites.
The presence of CYP52 members in selected yeast
species suggested that these P450s play a role in
Fig. 2. Comparative analysis of P450s in subphyla Saccharomycotina. A total of 172 P450s representing all P450
families found in Saccharomycotina were used for analysis. The P450 family, number of member P450s in a family
and percentage compared to overall P450 count (172 P450s) are also shown in the pie chart.
J PURE APPL MICROBIO, 8(SPL. EDN.), NOVEMBER 2014.
297KGOSIEMANG et al.:  P450OME OF SACCHAROMYCOTINA
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J PURE APPL MICROBIO, 8(SPL. EDN.), NOVEMBER 2014.
298 KGOSIEMANG et al.:  P450OME OF SACCHAROMYCOTINA
Table 3. General function and substrate specificity of fungal P450s. P450s that are functionally characterized
and present in Saccharomycotina species are shown in the table.
P450 General function Substrate Product(s) Reference
CYP51 Cell membrane Lanosterol 4,4-dimethylcholesta-8,14,24- 47
sterol trienol
biosynthesis 24-methylene-24,25- 4,4-dimethylfecosterol
dihydrolanosterol
CYP52 Alkanes and fatty Arachidonic acid 20-Hydroxy-5,8,11,14- 48
 acid icosatetraenoic acid
hydroxylation Dodecane 1-Dodecanol
2-Dodecanol
Hexadecane 1-Hexadecanol
2-Hexadecanol
Octadecane 1-Octadecanol
2-Octadecanol
Decane 1-Decanol
2-Decanol
Tetradecane 1-Tetradecano
2-Tetradecanol
Lauric acid 12-Hydroxydodecanoic acid 11-
Hydroxydodecanoic acid
Myristic acid 14-Hydroxytetradecanoic acid
13-Hydroxytetradecanoic acid
Palmitoleic acid 16-Hydroxy-9-hexadecenoic acid
 15-Hydroxy-9-hexadecenoic acid
Palmitic acid 16-Hydroxyhexadecanoic acid
15-Hydroxyhexadecanoic acid
alpha-Linoleic acid 18-hydroxy-alpha-linoleic acid
17-hydroxy-alpha-linoleic acid
Linoleic acid 18-hydroxy-linoleic acid
17-hydroxy-linoleic acid
Oleic acid 18-Hydroxy-9-octadecenoic acid
 17-Hydroxy-9-octadecenoic acid
Stearic acid 18-Hydroxyoctadecanoic acid
17-Hydroxyoctadecanoic acid
CYP56 Synthesis of component N-formyl tyrosine N,N’-bisformyl dityrosine 49
 of outer spore wall layer
CYP61 Cell membrane sterol Ergosta 5,7,24(28)-trienol Ergosta 5,7,22,24(28)-tetraenol 50
biosynthesis
CYP504 Phenylacetate Phenylacetate 2-Hydroxyphenylacetate 51
degradation
3-Hydroxyphenylacetate Homogentisate 52
3,4-Dihydroxyphenylacetate 2,4,5-Trihydroxyphenylacetate
the adaptation of yeast species to particular
ecological niches, e.g. utilization of specific
nutrients, such as alkanes, as carbon source19,20.
The distribution of other P450 families and the
number of member P450s across the selected yeast
species can be obtained in Table 2. One interesting
observation is that the CYP548 P450 family is
present only in Y. lipolytica.
The number of P450s in the yeast
genomes was lower compared to the fungal species
from other fungal phyla, Basidiomycota,
Zygomycota and Chytridiomycota, and even
compared to the subphylum Pezizomycotina12, with
few exceptions, suggesting the significant loss of
P450s in saccharomycotina species. The
phenomenon of the presence of a low number of
J PURE APPL MICROBIO, 8(SPL. EDN.), NOVEMBER 2014.
299KGOSIEMANG et al.:  P450OME OF SACCHAROMYCOTINA
P450s in Saccharomycotina species is mentioned
in earlier studies3,21,22. It has been shown that a
large number of P450s, especially certain P450
families that are highly enriched in Basidiomycota,
play a key role in the adaptation of Basidiomycetes
to different ecological niches, especially in wood
degradation, by performing extraordinary catalytic
activities5,7. This suggests that P450 numbers in
an organism can be directly linked to the
physiology/ecological niches of the organism.
Significant loss of P450s and the role of enrichment
of certain members of P450 families in few
Saccharomycotina species physiology are
discussed in the next section.
Role of P450s in Saccharomycetes physiology
Based on the literature available on the
functional characterization of P450s (Table 3) and
the characteristic life style of yeast species (Table
1), we conclude that Saccharomycotina species
retained P450s that are critical in their physiology.
For example, CYP51 and CYP61 P450s play a key
role in the synthesis of membrane components and
CYP56 is involved in the synthesis of the outer
spore wall layer (Table 3). CYP52 family members
that are enriched in Candida species and Y.
lipolytica are involved in organisms’ primary
(oxidation of fatty acids) and secondary (oxidation
of alkanes) metabolism. CYP52 P450s of C.
tropicalis play a key role in the production of
biotechnologically valuable long-chain
dicarboxylic acids23. The presence of the highest
number of CYP52 members in both yeast species
gives the species an advantage to utilize alkanes
as a carbon source (adaptation to different
ecological niches). A deletion mutant of Y.
lipolytica, where 12 CYP52 genes were deleted,
was unable to utilize n-alkanes (10-16 carbon
length) as a carbon source20. This strongly
supports the argument that P450s play a key role
in the adaptation of yeast species to diverse
ecological niches. CYP504 oxidizes simple aromatic
compounds such as phenylacetate and helps
organisms to grow on this organic compound.
Considering the functional properties of
characterized P450s, we conclude that the orphan
P450s in Saccharomycotina possibly play a role in
organisms’ physiology, either in synthesis of
cellular components or utilization of simpler organic
molecules.
CONCLUSIONS
The ecological niches of yeast species
are highly enriched with simpler organic nutrients,
unlike other fungal species such as Basidiomycetes
that degrade wood, a complex polymer, to gain
access to nutrients. Because of the complex
metabolic process involved in degradation of wood
components, Basidiomycetes are enriched with
catalytically diverse P450s5,7. This clearly suggests
that ecological niches of organisms play a critical
role in shaping their genome content. Further
evidence of this phenomenon can be obtained from
the fact that Cryptococcus neoformans and
Tremella mesentrica contain eight and 10 P450s in
their genomes compared to other basidiomycetes7.
C. neoformans is a well-known human pathogen
and T. mesenterica is a parasite fungus. It is shown
that these non-wood-degrading Basidiomycetes
lost P450s owing to their different ecological niches
(life style pattern) compared to the wood-
degrading basidiomycetes7. It has been suggested
that yeast-forming fungi (Saccharomycotina and
Taphynomycotina) have small P450omes, while
mycorrhizal relationships and complex nutrient
degradation seem to enhance P450
diversification21. Considering the type of ecological
niches adapted by yeast species, it is clear that
these species do not need a large number of P450s.
The presence of abundant and simpler organic
nutrients and adaptation of yeast species for
efficient utilization of these simpler nutrients further
compacted their genomes and they lost a
considerable number of P450s in their genome
during evolution.
ACKNOWLEDGMENTS
KS and SSM thank the Central University
of Technology (CUT), Bloemfontein, Free State for
a grant from the University Research and
Innovation Fund. SSM also thanks the National
Research Foundation (NRF), South Africa for the
research grant. The authors want to thank Dr David
R Nelson, University of Tennessee, USA for
naming the P450 family and subfamilies. The
authors also want to thank Ms Barbara Bradley,
Pretoria, South Africa for English language editing.
J PURE APPL MICROBIO, 8(SPL. EDN.), NOVEMBER 2014.
300 KGOSIEMANG et al.:  P450OME OF SACCHAROMYCOTINA
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