Lasbela, U. J.Sci.Techl., vol. V, pp.201-207, 2016
ISSN
2306-8256
REVIEW ARTICLE
Biotechnology of Penicillium Genus
Fatima Sharaf Ali¹, Khalid Mehmood1, Muhammad Anwar1, Ali Akbar2, Samiullah3, Junaid
Baber4, Said Qasim5, Imran Ali1,6*
1
Institute of Biochemistry, University of Balochistan, Quetta, Pakistan
2
Department of Microbiology, University of Balochistan, Quetta, Pakistan
3
Department of Chemistry, University of Balochistan, Quetta, Pakistan
4
Department of Computer Science, University of Balochistan, Quetta, Pakistan
5
Department of Geography, University of Balochistan, Quetta, Pakistan.
6
Plant Biomass Utilization Research Unit, Botany Department, Chulalongkorn University,
Bangkok, Thailand.
Abstract:- Penicillium play a vital role in biotechnology. There are over 300 species of Penicillium which
are beneficial to mankind from various features and many are still unexplored that need to be identified. Penicillium
exigency ranges from bioremediation, enzyme and biogas begetter, antioxidant source, vitamin producers, pigment
provenance for textile/cosmetics colorant and the most and eminent of all as antibiotic agents and some even with
antifungal properties. These tiny organisms are more than blessings for mankind. The human population is increasing with
increasing demand therefore there is prompt need of research in this trivialized branch of science.
Keywords: Fungi, Penicillium, Biotechnology. Dye.
grown in cheese and meat that is healthy for
mankind so the Penicillium and the trial to
its discoveries should be enhanced to
broader aspects (Ali et al., 2014a,b).
INTRODUCTION
Penicillium
lies in phylum
Ascomycota of kingdom fungi with the
greatest importance. Up to date over 300
species of Penicillium are discovered some
of which are Penicilium albicans,
Penicillium expansum, Penicillium notatum,
Penicillium purpurogenum, Penicillium
roqueforti,
Penicillium
restrictum,
Penicillium
glandicola,
Penicillium
digitatum,
Penicillium
flavigenum,
Penicillium inflatum, Penicillium tricolor,
Penicillium oxalicum, and Penicillium
viticola. After Penicillium, Aspergillus has
the second greatest importance in kingdom
fungi.
The importance of Penicillium is
due to its wide application in biotechnology
to benefit mankind. The low cost production
of valuable chemicals, enzymes, vitamins,
proteins and organic molecule has become
possible due to Penicillium. Baking bread,
production of low cost high effective
pigments, killing bacteria and making cheese
is due to the qualities of Penicillium,
however it also functions in mycotoxin
production and food spoilage but its
economic utility has more importance than
negative list. Even edible Penicillium are
The body type of Penicillium
is thallus (mycelium). The hyphae of the
Penicillium are multinucleated, colorless and
septate. Ascospores and conidiospores are
the two types of spores formed in
Penicillium asexually and sexually. The
conidiospores appear on conidiophores
which are then spread and the colony
broadens while the ascospores are formed by
the fusion of antheridium and archegonium.
Penicillium lives in wide range of habitat
from cold to hot environment but most
specifically soil fungi prefer moderate and
cool environment. Penicillium lives as
saprophytes on foods and fruits. It appears
on different substances including buildings,
ceilings and machineries. The spores of
Penicillium are present in the environment
always.
Role in Biotechnology
Penicillium is very crucial due to
wide application in biotechnology. It is used
for the production of many gases like
Hydrogen and Methane gas, bioremediation
of heavy metals, enzymes, vitamins, cheese,
alcohol, and pigment production. Its
antibacterial
production and
genetic
engineering is already known to everyone
*Corresponding Author: [email protected]
201
Bioremediation
Industrial activities are causing
various environmental pollution which is
responsible for increased organic and
inorganic xenobiotics in the ecosystem.
Heavy metals, poly cyclic aromatic
hydrocarbons (PAHs), Phenol and phenolic
compound. The inhalation or presence of
these substances in the environment is
hazardous to mankind and all ecosystem
members. Penicillium and some other fungi
are greatly used for the bioremediation of
such toxic chemicals. The bioremediation
technologies used nowadays involve
physical, chemical and biological processes.
The biological process needs Penicillium
and some other fungi for the removal of
such xenobiotics from water, soil and air.
Fungi grow in aquatic sediments, terrestrial
habitats and water surfaces. They are also
important for natural remediation of the
environment. Fungi are beneficial over
bacteria in this sense as fungal hyphae can
penetrate into the contaminated soil for
thebioremediation action. Among all fungi,
Penicillium have received the maximum
attention in bioremediation. They have
shown their capability to deteriorate and
degrade different xenobiotic which are
termed
mycodeterioration,
and
mycodegradation respectively. Penicillium
simplicissimum YK is capable of degrading
polyethylene with molecular weight (MW)
of 400 to 28000 (Onodera et al.,
2001).Moreover fungi have the ability to
produce extracellular enzymes for the
degradation and assimilation of pollutants.
Most of Penicillium strains are documented
to be halotolerant, some halophile, and some
as obligate halophile. The halophilic
Penicillium strains have better tendency of
bioremediation as the factories waste contain
high concentration of salt with heavy
metals, organic and inorganic acids and
radionuclides therefore the capacity of
halophilic fungi to remediate pollutants in
the presence o f salt becomes e a s i e r . P.
janthinellum F-13 has the ability to
minimize the aluminum toxicity and
production of citric acid (Zhang et al.,
2002). Recently Fan (2008) reported the
removal of Zn, Cd and Pb by P.
simplicissimum and this fungus has very high
potential for bioremediation of metals.
Penicillium not only helps in bioremediation
but also helps in accumulation of some
metals like uranium.
P. digitatum
accumulates uranium uranyl chloride
aqueous solution. The fungus was treated in
boiling water and alcohol, it increased the
uptake ability to 10,000 parts per million
(dry weight) moreover the mycelium of this
Penicillium has great tendency to bind with
Pb. Zn, Ni and Cd (Galun et al., 1983;
Mendil et al., 2008). Say R (2003) reported
P. canescence capable of removing As, Hg,
Cd and Pb ions from the aqueous solution of
biosorption.
He
also
reported
P.
purpurogenum to remediate high amount of
Cr. The Cr absorption ability tends to
increase in the first four hours and the
level turns to equilibrium.
Penicillium
strains
not
only
remediate heavy metals, they are also widely
used
to
remediate
PAHs.
These
hydrocarbons exist everywhere in the world
but due to different strains of Penicillium this
has become possible to remediate such
PAHs. Many of the techniques used now
days for remediation of PAHs include
biological processes.
The efficiency of
removal depends on type and concentration
of PAH, moisture content, temperature, soil
characteristics, microorganism, and oxygen
concentration. P. frequentans has the ability
to
remove
phenanthrene.
Oxygen
concentration has a substantial effect in this
action (Estrada et al., 2006). Fluorine
degradation in the presence of cyclodextrin
was documented (Garon et al., 2004).
Pyrene is a four ring structure known as
genotoxic and it indicates PAH contaminated
wastes
working as an indicator. P.
janthinellum SFU403 has the ability to
remove Pyrene. This strain of Penicillium
was obtained from petroleum contaminated
soil (Boonchan et al., 2000) while
Saraswathy and Hallberg (2000) reported for
the first time that fungi uses pyrene as the
energy source and sole carbon in liquid
202
culture.
utilized in the production of other substances
or other purposes.
For enzyme production, filamentous
fungi are excellent source. They are already
famous due to production of extracellular
protein. Cellulolytic enzymes are produced
by Trichoderma reesei, a fungus of
ascomycetes. Cellulase is secreted in a large
amount which can benefit industries
(Kubicek et al., 2009). This fungus is known
as efficient producer of extracellular
cellulolytic enzymes (Juhasz et al., 2005;
Sehnem et al., 2006). P. chrysogenum also
produces extracellular protein with cellulase
activity. The same fungus when cultured on
media containing sawdust, wheat bran,
sugarcane pulp and oat spelt produces xylan.
P. purpurogenum is reported for the
production of extracellular xylanases
moreover xylanase A and xylanase B have
been purified and characterized from this
fungus (Belancic et al., 1995). Kunitz (1938)
reported a Penicillium sp. for the production
of kinase that synthesized trypsin from
trypsinogen. Penicillium oxalicum stated for
the production of caseinase enzymes with
milk clotting activity. Concisely there are
number of fungi that produce useful enzymes
that can benefit mankind.
Antibacterial and Antioxidants
Fungi are also useful for the removal
of olive mill wastewater (OMW). Olives
grown
in
Mediterranean
countries
comprises about 98% of global olive
production. Large quantity of OMW is
produced while olive oil production. 2.5
litre of waste is releases per litre of oil
produced. Methods of removal include
conventional, electrolysis, and aerobic
method while aerobic method is the best
method for removal of this waste water as it
contains phenolic, tannin and lignin
compounds. Penicillium produces such
enzymes which are very effective in
detoxification of this waste water.
Penicillium P4 when cultivated with OMW
caused phenolic reduction up to 54% and
COD reduction up to 61 % (Robles et al.,
2000).
Ethanol is produced by fermentation
distillation on a large scale in Europe and
USA. As a result of ethanol production high
amount of liquid waste is generated called
vinasses. About 9 to 14 L of waste water is
produced per liter of ethanol production. And
this waste water is highly acidic (pH 4-5)
with high organic content (COD 50 to
100 g/L) (Jiménez et al., 2006).
P.
decumbens was reported to degrade the
phenolic compounds of vinasse in batch
regime. About 74 % of phenol removal was
noted after 3 days of treatment (Jiménez et
al., 2005). P. decumbens also becomes
successful in decolorization of vinasse. 41 %
color decolorization was noted on the 5 th
day. Concisely, fungi play a vital role in
bioremediation of all type of pollutant
moreover i t is a cheap source for toxic
pollutant removal. It is also a natural
environment bioremediating agent. Currently
many halophilic species are identified that
had potential to remediate NaCl from its
medium (Ali et al., 2014).
The antibacterial activity of fungi is
already known to everyone. Fungi
specifically strains of Penicillium are having
tremendous
potential
of
producing
antibacterial agent and antioxidants. P.
chrysogenum has gained scientist’s attention
for production of antibiotic Penicillin. The
discovery of Penicillin goes back to
Alexander Fleming in 1928. Alexander told
his Penicillium as P. chrysogenum but
Houbraken et
al. (2011) declared
Alexander’s Penicillium as P. rubens. These
two species are phenotypically similar but
extrolite analysis demonstrate that secalonic
acid D and F or a metabolite related to
lumpidin
are
produced
by
P.
chrysogenum while such metabolites are not
produced by P. ruben (Houbraken et al.,
2011). Antibiotics from P turbatum broth
fermentation were obtained by Michel and
Penicillium as Enzyme Producer
There are number of Penicillium that
produces extracellular enzymes in their
culture media which are very useful to
mankind. These enzymes are collected and
203
his colleagues (2006). Currently in 2006 a
new antibiotic leucinostatin was discovered
and identified by Arai and his friends from
P. lilacinum. It was obtained in the form of
white prisms.
Microorganism typically
bacteria has the habit of going under
mutation or adopting resistance against
antibiotics therefore, there is continuously
need of research on antibiotics and
discoveries of new antibiotics from new
species.
many contributions to food science and food
industries (Khachatourians
and
Arora,
2001). Other members of fungi are already
known as food consumption and application
at food industried like Yeasts, Candida
albicans, Pseudotropicalis, Pachysolen
tannophilus,
Phaffia
rhodozyme,
Saccharomyces
cerevisiae,
Yarrowia
lipolytica, Pichia guilliermondii and
Penicillium is on the importan lines of
description as food consumption and
applications but for any microorganism to be
used in food industry must have generally
recognized as safe (GRAS) status according
to FDA (Food and Drug Administration).
Vitamins, organic acids, organic
molecules, pigments, volatile and nonvolatile
flavors are important compounds in food
biotechnology. Their production methods
rely on microorganism specifically fungi but
for any microorganism Specific enzymes
produced by fungi contribute value in food
biotechnology. The global market demand
of enzymes comes from fungi and bacteria.
P. roqueforti is an example of edible
Penicillium. It is a common fungus
widespread in the nature, used in the
production of blue cheese. The consumption
of blue cheese is documented from AD 50
(Ridgway, 2004). In short fungi have a great
impact on food science from baking bread at
home till the large scale production
industries.
Penicillium and Cancer
However Penicillium itself is
responsible for causing different diseases in
living being but its services in disease
treatment like Cancer is reported in the
recent years which is a massive achievement
in disease treatment. Species of Penicillium
are being used in nanotechnology due to its
cytotoxic effect against cancer. Mishra and
collogues
recently
have
used
P.
brevicompactum biomass for nanoparticle
formation against cancer. She stated that the
species has cytotoxic effect against mouse
mayo blast cancer C2C12 cells (Mishra et
al., 2011). 3-O-methylfunicone produced
from P. pinophilum can inhibit stem cells of
breast cancer stated by Buommino and his
Ali et al. (2014a,b) have declared
numbers
of
filamentous
fungi
as
antioxidants and antibacterial producer.
Antioxidant 2, 3-dihydroxy benzoic acid was
isolated from P. roquefortii (Hayashi et al.,
2014). P. citrinum was reported positive for
the production of novel type of antioxidants
in its culture broth. Concisely Penicillium
and the services it provides to mankind are
unlimited, many more are still undiscovered
which need to be discovered, identified and
modified for biotechnological applications.
Biogas
Energy has become an essential part
of our daily life. Our maximum necessities
are being fulfilled by fossil fuels. Due to
excessive use, fossil fuels are decreasing.
They are also reported as one of the reason
of increasing global warming and acid rain
therefore there is need of another source of
energy (Das and Vezirolu et al., 2011).
Scientist has already discovered many other
ways of energy production in which
mycologists are in the line of description
after solar energy.
Biogas and bio power are being synthesized
and used currently in number of countries by
waste water and renewable resources in
which microorganism play a vital role.
Presently many countries are having biogas
plants. Biogas from Chlorella sp. (Ali et al.,
2011), cow dung, and fruit peels are already
reported and species of Penicillium are
treated with the substrate to enhance biogas
production. Species of Penicillium are under
research for biogas production (Xiao-ming et
al., 2008).
Penicillium and Food
Fungal biotechnology has
done
204
friends (2011).
Penicillium and Pigment
There are number of Penicillium
species that secrete pigment in its media
which are dried and used as colorant in food,
cosmetics, textile and pharmaceutical
industries. Color upholds a vital role with the
food we consume. It is the primary
identification o f a consumer for buying
food. At markets purchaser are attracted
toward good colored products and avoid
synthetic dyed foods. We purchase those
products that are felt well by our eyes.
Pigment from P. oxalicum, arpink red color
is being used extensively in food industries
(Dufossé, 2006). Furthermore the food
industries are cracking to use natural dyes
substituting synthetic dyes using plant and
microbial pigments. A blue pigment is
reported from P. herquei (Frank et al.,
1951). The demand of natural colorant in
food and cosmetics have increased that
people are ready to pay a premium. Stich
(2002) claimed exquisitely that we eat with
our eye.
identified extremophiles P. species must be
enhanced to reach a milestone.
REFERENCES
Ali, I.,
Akbar,
A., Anwar, M., Yanwisetpakdee,
B., Prasongsuk, S., Lotrakul, P. and
Punnapayak, H., 2014a. Purification and
characterization
of
extracellular,
polyextremophilic a- amylase obtained from
halophilic Engyodontium album. Iranian J.
Biotech., 12: 35-40.
Ali, I.,
Siwarungson,
N.,
Punnapayak,
H.,
Lotrakul, P., Prasongsuk, S., Bankeeree,
W. and Sudip, K., Rakshit., 2014b.
Screening Of Potential Biotechnological
Applications From Obligate Halophilic
Fungi, Isolated From A Man-Made Solar
Saltern Located In Phetchaburi Province,
Thailand. Pak. J. Bot., 46(3): 983-988.
Ali, I., Rakshit, S.K. and Kanhayuwa L., 2011.
Biohydrogen
production
from
microalgae of Chlorella sp. The International
Conference on Sustainable Community
Development, Khonkaen Thailand,
74-77.
Ali, I., Akbar, A., Yanwisetpakdee, B., Prasongsuk , S.,
Lotrakul, P. and Punnapayak, H., 2014.
Purification, characterization and potential of
saline waste water remediation of a
polyextremophilic
a-amylase
from an
obligate halophilic Aspergillus gracilis
Biomed Research International Volume
2014,
Article ID 106937, 7 pages,
http://dx.doi.org/10.1155/2014/106937.
Arai T., Mikami, Y., Fukushima, K., Utsumi, T. and
Yazawa, K.,2006. A new antibiotic,
leucinostatin, derived from Penicillium
lilacinum.
The
Journal
of
Antibiotics,http://doi.org/10.7164/antibiotics.
26.
157.
Belancic, A., Scarpa, J., Peirano, A., Díaz, R., Steiner,
J. and Eyzaguirre, J. 1995. Penicillium
purpurogenum produces several xylanases:
Purification and properties of two of the
enzymes. Elsevier, Journal of Biotechnology.
Doi: 10.1016/0168-1656(95)00057- W.
Boonchan, S., Britz, M.L. and Stanley, G.A.,
2000.Degradation and mineralization of
high-molecular-weight polycyclic aromatic
hydrocarbons by defined fungal-bacterial
cocultures.
Appl.
Environ.
Microbiol. 66:1007–1019.
Buommino, E., Tirino V., De Filippis, A., Silvestri, F.,
Nicoletti, R., Ciavatta, M. L., Pirozzi,
G.,Chinedu N. S, Okochi, VI, Smith, H.A.,
Okafor,
U.A., Onyegeme-Okerenta, B.M.
and Omidiji, O., 2007. Effect of carbon
sources on cellulase (EC3.2.1.4) production
by Penicillium chrysogenum PCL501. Afr.J.
Mycol Biotechnol, 1:006–010.
CONCLUSION
Species
of
Penicillium,
are
extensively important in biotechnology to
benefit mankind and its environment. The
importance of any microorganism can be
enumerated by estimating its negative list
with the positive. Species of Penicillium
therefore occupy the most important place in
biotechnology. Scientists can not consider
development in human life without the aid
of Penicillium. In the current years obligate
and extreme halophilic, thermophile,
acidophilic and basophilic Penicillium are
identified from the extreme environments of
the world which can come with huge
achievement after research. As the
extreme halophilic Penicillium is reported
for scavenging salt from its medium so it is
potential to desalinize sea water. The global
warming is one of the biggest problem for
the scientist round the globe, the extreme
thermophilic Penicillium are potential to
minimize global heat.
The research and
trials on Penicillium Species and the newly
205
Das, D. and Vezirolu, T., N.2001. Hydrogen
production by biological process: a survey of
literature. International Journal of Hydrogen
Energy. pp. 13-28.
Dufossé, L., 2006. Microbial Production of Food
Grade Pigments. Food Grade Pigments, Food
Technol. Biotechnol.44 (3): 313–321. ISSN
1330-9862.
Schmoll, M. and Seiboth, B.,
2009.
Metabolic engineering strategies for the
improvement of cellulose production by
Hypocrea jecorina. Biotechnol Biofuels,
2:19. doi: 10.1186/1754-6834-2-19.
Kunitz , M., 1938. Formation of trypsin from
trypsinogen by an enzyme produced by a
mold of the genus Penicillium.The journal
of general physiology. 21(5): 601-620 Doi
http://dx.doi.org/10.1085/jgp.21. 5.601.
Mendil, D., Tuzen, M. and Soylak, M., 2008. A
biosorption system for metal ions on
Penicillium italicum-loaded on Sepabeads SP
70 prior to flame atomic absorption
spectrometric determinations. J. Hazard.
Mater.152:1171–1178.
Michel, K. H., Chaney, M. O., Jones, N. D.,
Hoehn, M. M. and Nagarajan, R., 2 0 0 6 .
Epipolythiopiperazinedione
antibiotics
fromPenicillium
turbatum, The Journal of Antibiotics.
http://doi.org/10.7164/antibiotics.27.57.
Estrada, M.J., Amezcua-Allieri, M.A., Alvarez, P.J.J.,
Rodríguez-Vázquez, R., 2006. Phenanthrene
removal by Penicillium frequentans grown
on a solid-state culture: effect of oxygen
concentration.
Environ.
Technol.,
27:1073–1080.
Fan, T., Liu Y., Feng, B., Zeng, G., Yang, C., Zhou,
M., Zhou, H., Tan, Z. and Wang, X., 2008.
Biosorption of cadmium (II), zinc (II) and
lead (II) by Penicillium simplicissimum:
Isotherms, kinetics and thermodynamics. J.
Hazard. Mater. 160:655–661.
Galun, M., Keller, P., Malki, D., Feldstein, H., Galun,
E., Siege,l SM. and Siegel, B.Z., 1983.
Removal of uranium (VI) from solution by
fungal biomass and fungal wall-related
biopolymers. Science, 219:285–286.
Generally
Mishra, A., Tripathy, S. K., Wahab, R., Jeong, S-H.,
Hwang, I., Yang, Y-B., Kim, Y. S., Shin,
H. S. and Yun,
S.I., 2011. Microbial
synthesis of gold nanoparticles using the
fungus Penicillium brevicompactum and
their cytotoxic effects against mouse mayo
blast cancer C2C12 cells. Appl Microbiol
Biotechnol
(2011)
92: 617. Doi:
10.1007/s00253-011-3556-0.
Onodera, Y.KH., Mukumoto, Y., Katsuyaya, Y.,
Saiganji, A., Tani, Y., 2001. Degradation
of polyethylene by a fungus, Penicillium
simplicissimum YK. Polym. Degrad. Stabil.
72:323–
327.
Ridgway, J., 12 October 2004. The Cheese
Companion. Running Press. p. 10. ISBN
978-0-7624-1956-2. Retrieved, 2 February
2013.
Robles, A., Lucas, R., de Cienfuegos, G.A. and
Galvez, A., 2000. Biomass production and
detoxification of wastewaters from the olive
oil industry by strains of Penicillium isolated
from wastewater disposal ponds. Biores.
Tecnhol. 74:217–221.
Saraswathy, A.,
Hallberg, R., 2002. Degradation of
pyrene by indigineous fungi from a former
gasworks site.FEMS Microbiol. Lett.
210:227–232.
Say, R., Yilmaz,
N. and
Denizli,
A.,
2003. Removal of heavy metalions using the
fungus Penicillium canescens. Adsorpt. Sci.
Technol. 21:643–650.
Sehnem , N.T.,
de Bittencourt, L.R., Camassola,
M., Dillon, A.J.P., 2006. Cellulase
production by Penicillium echinulatum on
lactose. Appl Microbiol Biotechnol. 72:163–
167. doi: 10.1007/s00253-005-0251-z.
Stich, E., Chaundry, Y. and Schnitter, C., 2002.
Recognized
as
Safe
(GRAS).
http://www.fda.gov/.Retrieved 30/10/2016.
Hayashi, k. I., Suzuki, K., Kawaguchi, M., Nakajima,
T., Suzuki, T., Numata, M., and Nakamura,
T., 2014. Isolation of an Antioxidant from
Penicillium roquefortii IFO 5956, Journal
Bioscience,
Biotechnology,
and
Biochemistry,
Volume
59,
1995.
http://dx.doi.org/10.1271/bbb.59.31.
Houbraken, Jos; Frisvad, Jens C.; Samson', Robert A.
2011. Fleming's Penicillin producing strain is
not Penicillium chrysogenum
but
P.
rubens.
ISSN-2210-6340 (Print); ISSN
2210-6359.
Jimenez, A.M., Borja, R., Martin, A., Raposo, F.,
2005. Mathematical modelling of aerobic
degradation of vinasses with Penicillium
decumbens. Process Biochem, 40: 2805–281.
Jiménez, A.M., Borja, R., Martín, A. and Raposo, F.,
2006. Kinetic analysis of the anaerobic
digestion of untreated vinasses
and
vinasses
previously treated with
Penicillium decumbens. J. Environ. Manage,
80:303–310.
Juhasz, T., Szengyel, Z., Reczey, K., Siika- Aho, M.
and Viikari, L., 2005. Characterization of
cellulases and hemicellulases produced by
Trichoderma reesei on various carbon
sources. Process Biochem, 40:3519–3525.
doi:10.1016/j.procbio.2005.03.057.
Khachatourians, G G. and Aroram D.K., 2001.
Applied Mycology and Biotechnology
Volume 1: Agriculture and Food Production.
Amsterdam: Elsevier Science.
Kubicek, C.P.,
Mikus,
M.,
Schuster,
A.,
206
Colour you eat with your eyes, Int. Food
Ingred. 1: 6–8.
Tufano M. A. 3-O-methylfunicone, from Penicillium
pinophilum, is a selective inhibitor of breast
cancer stem cells in Cell prolieferation.
DOI: 10.1111/j.1365- 2184.2011.00766.x.
Zhang, D., Duine, J.A. and Kawai, F., 2002. The
extremely high Al resistance of Penicillium
janthinellum F-13 is not caused by internal
or external sequestration of Al. Biometals.
15:167–174.
Received September 10th, Accepted December 20th
2016
Manuscript can be viewed online at:
http://www.lujst.com/
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