66
The genus Chrysosporium,
its physiology and biotechnological
potential
Rajendra Kumar Singh Kushwaha
Department of Botany, Christ Church College, Kanpur, India
Summary
Key words
The genus Chrysosporium is reviewed including its 11 species without intercalary conidia, 29 species with intercalary conidia, three species with uncertain
position and four undescribed species with their report of isolation, physiology,
antagonistic, keratinolytic and pathogenic potentials. Research revealed considerable biotechnological potential for recycling keratinous waste in soil and
secretion of enzymes and antimicrobials. Its taxonomic relation with dermatophytes and their relatives is of immense importance. Germplasm collection of
Chrysosporium and its teleomorphic connections is aplaused.
Chrysosporium, Biotechnological Potential, Review
The genus Chrysosporium was introduced by
Corda [1]. Saccordo [2] placed it in synonymy with
Sporotrichum. Later this genus was reviewed [3-5].
Twenty two species of Chrysosporium were recognised
[5]. Since then several species have been added to this
genus. Large number of workers have isolated
Chrysosporium species from many different habitats
around the world. Until recently, interest was restricted
only to reports of its occurrence but as research into this
fungus has gained momentum it has been evaluated as a
potential fungus. Recently the research carried out on
Chrysosporium has increased markedly and information
has begun to appear. While isolating Chrysosporium by
hair baiting and other methods its potential to degrade
keratin was particularly emphasized. In view of this, its
activity in soil and water sediments of polluted and fresh
water sites is also receiving attention. Perhaps some species of Chrysosporium may be utilized for recycling of
keratinous waste in soil and as water pollution indicators
which certainly pave the way towards a congenial environment. Secretion of some of their metabolites, particularly enzymes and antimicrobials, is gaining the attention
of pharmaceutical industries. The resemblance of
Chrysosporium to dermatophytes, and their pathogenic
potential, is directly related to health of human beings and
animals. In addition to the keratinous substrates, this
genus is now being found associated with other non keratinous substrate too. This potential, and its similarities to
Myceliophthora, Emmonsia, Zymonema, Geomyces,
Corresponding address:
Dr. Rajendra K.S. Kushwaha
Department of Botany,
Christ Church College,
Kanpur 208 001, India
Tel: +91 512 637 318; Fax +91 512 311 627
©2000 Revista Iberoamericana de Micología
Apdo. 699, E-48080 Bilbao (Spain)
Trichosporiella,
Malbranchea,
Ovadendron,
Botryotrichum, Sepedonium, Mycogone, Sporotrichum
and others and associated teleomorphs, is of immense
diagnostic importance. Long term studies [6-11] on the
biology of Chrysosporium, and other scattered reports,
revealed that its wide distribution is due to its antagonistic
potential and ability to produce enzymes and other extracellular metabolites [12-14].
Looking into all the above potential of
Chrysosporium, it was intended to review all possible
available work on this genus, which has not been reviewed before, in spite of the fact that the amount of new
information has grown enormously in recent times.
Forty - seven species of Chrysosporium are listed
here along with their report of isolation including some
salient features.
Species with intercalary conidia
White colony
1.Chrysosporium anamorph of Rollandina vriesii
Apinis Trans. Brit. Mycol. Soc. 55:501, 1970.
25-30 mm on PYE in 14 days, terminal and lateral
conidia smooth and thin walled, 1 celled, 3-6x2-3 µm,
wide scar, 1-2 µm, keratinolytic.
2.Chrysosporium anamorph of Arthroderma curreyi Berk Outl. Bri. Fungol. 357, 1860.
30-50 mm on SGA in 14 days, terminal and lateral
conidia mostly sessile, smooth or slightly echinulate, thin,
1 celled, rarely 2 celled, 3-6x2-3 µm, wide scar, 1-2 µm,
keratinolytic [15-19].
3.Chrysosporium anamorph of Arthroderma cuniculi Dawson Sabouraudia 2:187, 1963.
35-60 mm on SGA in 14 days, terminal and lateral
conidia sessile, smooth, thin walled, 1 celled or 2-3 celled,
mostly 3-8x2-3 µm, scar 1-2 µm, keratinolytic [16,17,20].
4.Chrysosporium anamorph of Pectinotrichum llanense Varsavsky & Orr Mycopath. Mycol. Appl. 43:231,
1971.
5-15 mm on Czapeck agar in 14 days, terminal and
lateral smooth and thin walled, 1 celled, 4-6.5x2-3 µm,
scar 1-1.5 µm, keratinolytic [21].
Chrysosporium and its potential
Kushwaha RKS
5.C. synchronum van Oorschot Stud. Mycol.
20:42, 1980.
80 mm on PYE in 7 days, terminal and lateral
conidia thin walled, smooth or echinulate, 1 celled,
7.5-11x4-5.5 µm, scar 0.5-1 µm, not keratinolytic.
Other than white
6.C. sulfureum (Fiedl.) van Oorschot & Samson
Stud. Mycol.20:28, 1980.
10-15 mm on PYE in 14 days, pale creamy yellow,
terminal and lateral mostly sessile, 2–8 per conidiogenous,
smooth and thin becoming thick walled, some times echinulate, 1 celled, 3-8x3-6 µm, wide scar, 1.4 µm, not keratinolytic, shows preference for fatty or calcium rich
material [22-24].
7.C. georgii (Varsavsky & Ajello) van Oorschot
Stud. Mycol.20:31, 1980.
10-30 mm on 2% malt agar in 14 days, white or
pale buff or pink, terminal and lateral smooth and thin
walled, 1 celled, rarely 2-3 celled, 3-8x2-3 µm, wide scar,
0.5-4.5 µm, keratinolytic [25,26].
8.C. lucknowense Garg Mycopath. Mycol. Appl.
30: 224, 1966.
55 mm on SGA in 14 days, cream, terminal and
lateral 1-4 conidia on one hyphal cell in close proximity,
thin, 1 celled, 2.5-11x1.5-6 µ m, keratinolytic
[22,23,26,27].
9.C. filiforme Sigler, Carmichael & Whitney
Mycotaxon 14:261, 1982.
40 mm in 21 days on PYE, white to buff, terminal
or lateral conidia 0-1 or rarely 2 septate, sessile or on
short pedicell, smooth, filiform upto 40 µm long, not keratinolytic.
10. C. mephiticum Sigler Can. J. Bot. 64: 1212,
1986.
50-62 mm on PYE and CER in 25 days, creamy
white, terminal and lateral smooth, mostly sessile, in close
proximity, 2.5-3.5x2.5-3 µm, scar 1-1.5, keratinolytic,
strong and pungent odour.
11.C. gourii Jain, Deshmukh & Agrawal Mycoses
36:77, 1993.
74 mm on SGA in 18 days, white to cream to
yellowish brown, terminal and lateral, smooth or rough,
thin walled, 1 celled, 2.5-6x2-4 µm, keratinolytic.
Species with intercalary conidia
White colony
12.C. queenslandicum Apinis & Rees Trans. Brit.
Mycol. Soc. 67:524,1976.
55-65 mm on PYE in 14 days, white, intercalary
conidia several, smooth and slightly thick, 1 celled, 3.59.5x3.6 µm, broad basal scar, keratinolytic [17,21-23,25-27].
13.Chrysosporium anamorph of Gymnoascus
demonbreunii Ajello & Cheng Mycologia 59:692,1976.
25-33 mm in 14 days on hay infusion agar, white,
intercalary conidia most abundant, smooth and thin,
5-11x4.5-6 µm, conidia always terminal, never lateral,
smooth, thin walled, 1 celled, 6-9x5-6 µm wide basal scar,
1.5-2.5 µm, keratinolytic [21].
14.C. carmichaelii van Oorschot Stud. Mycol.
20:15,1980.
15-30 mm in 14 days, white, intercalary less abundant, smooth, thin, 3-6x1.5-3 µm, terminal and lateral
smooth or rarely rough, 1 or rarely 2 celled, 3-6x3-3.5
µm, not keratinolytic [17,18,22,23,27,28].
15.C. tropicum Carmichael Can. J. Bot. 40: 1170,
1962.
50-60 mm in 40 days, white, intercalary not com-
67
mon, smooth and slightly thick, 3-4x6-10.5 µm, terminal
and lateral smooth or slightly thick, 1 or rarely two celled,
3.5-7.5x3-4.5 µm, basal scar 1.5-2 µm, keratinolytic [1618,21-23,25-33,36-53].
16.C. xerophilum Pitt Trans. Brit. Mycol. Soc. 49:
468, 1966.
85 mm on cherry deccoction agar in 7 days, white,
intercalary conidia not always abundant, sometimes in
chains of 2-6, smooth and thin or thick walled, 7.5-12x34.5 µm, terminal and lateral, smooth, thin or thick walled,
4-13x3-10 µm, wide basal scar, 1.5 µm, weak keratinolytic, osmophilic [21].
17.C. pannicola (Corda) van Oorschot and Stalpers
Stud. Mycol. 20:43,1980.
20-38 mm on PYE in 14 days, white, intercalary
less abundant, smooth or echinulate, thick walled, 4-8x2-3
µm, keratinolytic [15,17,18,20,22,23,26,27,34,54,55].
18.C. indicum (Randhawa & Sandhu) Garg
Sabouraudia 4:262,1966.
40-50 mm on PYE in 14 days, white, intercalary
conidia less abundant, smooth or slightly echinulate, thin
walled, 6-12x2-3.5 µ m, keratinolytic [15-23,25-32,40,
44,45,48,54, 56].
19.C. sinense Liang Acta Mycologica Sinica
10:50,1991.
30 mm in 20 days at 18°C on SDA, intercalary
conidia abundant, 1.2-3.7x3.7-10 µm, terminal and lateral,
2.4-3.8x4.0-5.7 µm, development of synnemata, isolated
from endosclerotium of Cordyceps sinensis [Berk.] Sacc.
20.C. geophilum Kushwaha & Shrivastava Curr.
Sci. 58:970,1989.
60-70 mm on SGA, white, reverse pale creamy
brown, intercalary conidia less, rough or smooth walled,
2-4 µm, lateral conidia sessile or on short protrusions, initially echinulate becoming smooth on transfers, thick, 1 or
rarely 2 celled, 4-20x2-4 µm, keratinolytic.
21.C. botryoides Skou Mycotaxon 43:237,1992.
White colony, ramose on MGYA, conidia 1 celled,
thick walled, occur so close together that they look like
bunches of grapes, globose to pyriform, up to 10.2 µm,
intercalary conidia sparsely present, rounded in agar, single or a few in chains, not keratinolytic, osmophilic.
22.C. globiferum Skou Mycotaxon 43:237,1992.
White, dense with slightly ramose margin on
MGYA, conidia 1 celled, thick walled, globose to pyriform, upto 11.1 µm, intercalary conidia abundant, rounded in agar, not keratinolytic, osmophilic.
23.C. hispanicum Skou Mycotaxon 43:237,1992.
White, dense on MGYA, conidia 1 celled, thick
walled, globose to pyriform, upto 13.4 µm, in agar only
terminal conidia, intercalary conidia very sparcely in agar,
not keratinolytic, osmophilic.
24.C. holmii Skou Mycotaxon 43:237,1992.
White, intercalary conidia turncate never in pronounced amount, not in agar, terminal globose conidia,
pyriform up to 12.8 µm, not keratinolytic, osmophilic.
25.C. medium Skou Mycotaxon 43:237,1992.
White, flat on MGYA, conidia 1 celled, globose to
pyriform, upto 9.8 µm, intercalary abundant in agar, not
keratinolytic, osmophilic.
26.C. minor Skou Mycotaxon 43:237,1992.
White, flat, conidia 1 celled, globose to pyriform,
up to 8.4 µm, intercalary conidia single and rounded in
agar, not keratinolytic, osmophilic.
27.C. pyriformis Skou Mycotaxon 43:237,1992.
White, dense, flat on MGYA, conidia 1 celled,
thick walled, globose to pyriform up to 11.0 µm, intercalary conidia in agar more or less rounded, not keratinolytic, osmophilic.
68
Other than white
28.C. pseudomerdarium van Oorschot Stud.
Mycol. 20:14,1980.
7-20 mm on PYE in 14 days, white, locally pale
yellow, intercalary conidia intially smooth and thin walled
or becoming echinulate and / or thick walled, 3-6x3-6
µm, terminal and lateral in chains up to 4, initially smooth
and thin becoming echinulate, 1 celled, 2-6.5x1.5-5 µm,
weak keratinolytic [25,26].
29.C. merdarium (Link ex Grev.) Carmichael Can.
J. Bot.40:1160,1962.
30-35 mm on PYE in 14 days, white becoming
bright yellow, pink or green, intercalary conidia few, 512x3-6 µm, terminal and lateral 1 celled, 4-10x3-6 µm,
not keratinolytic. G. uncinatum do not develop conidia on
PYE while these are abundant on hay infusion agar
[15,16,20-24,31,36].
30.C. keratinophilum D. Frey ex Carmichael Can.
J. Bot. 40:1157,1962.
40-50 mm on PYE in 7 days, white to cream, sulphor yellow, intercalary conidia less abundant or rare,
smooth or echinate, thick walled, 6-25x3.5-7 µm, terminal
and lateral smooth or echinate, thick, 1 celled, 3.5-22x3.511 µ m, keratinolytic [15,17,20-23,25,26,30,32,34-38,
40,41,43-45,47,54-66].
31.C. inops Carmichael
Can. J. Bot.
40:1156,1962.
0.5-10 mm on PYE in 14 days, cream, conidia terminal or intercalary, smooth and thick walled, 1 celled,
6.5-12x5-9 µm, not keratinolytic.
32.Chrysosporium anamorph of Renispora flavissima Sigler et al Mycotaxon 10:133, 1979.
30-40 mm on PYE in 14 days, pale yellow, buff
centre, intercalary conidia rare, initially smooth, thin or
thick walled, verrucose, 1 celled, 6-8x5-8 µm, keratinolytic.
33.C. lobatum Scharapov Nov. Syst. niz. Rast.
15:144, 1978.
30-35 mm on PYE in 14 days, white becoming
pale green or pale grey, intercalary conidia rare, smooth,
thin walled becoming echinulate, thick walled, 1 celled, 34x2-3 µm, terminal and lateral, smooth and thin walled
becoming reddish brown to dark brown, echinulate, thick
walled 1 celled, 2-4x1.5-3.5 µm, scar 0.5-1.5 µm, keratinolytic.
34.C. vespertilium Guarro, Vidal & de Vroey
Mycotaxon 59:189,1996.
34-45 mm on PYE in 14 days, yellow, intercalary
conidia rare, smooth and thin walled, 1-3 celled, 5-20x2-5
µm, scar 2.5 µm, keratinolytic, coiled sterile hyphae.
35.C. pilosum Gene, Guarro & Ulfig Mycotaxon
50:107,1994.
Restricted, 0.4-0.7 mm on PYE, raised, yellowish
white to mustared yellow or light brown, reverse brownish
or dark brown, intercalary conidia smooth, thin becoming
thick walled and verrucose, 1 celled, 3.5-5.5x3-4 µm, terminal and lateral smooth and thick walled, becoming
coarsly verrucose, 1 celled, 4-6x3.5-5.5 µm, scar 1.5-2
µm, poor keratinolytic, broad, simple, thick walled, brownish sterile hyphae present.
36.C. europae Sigler, Guarro & Punsola Can. J.
Bot. 64:1212,1986.
50-60 mm on PYE in 35 days, vinaceous buff or
brown diffusing pigment, intercalary 4.5 µm, terminal
and lateral 8.5x2.5-3.5 µm, keratinolytic.
37.C. zonatum Al - Musallam & Tan Persoonia
14:69,1989.
55 mm on PYE in 14 days, white to buff, intercalary conidia abundant, 5-12x2-4 µm, terminal and lateral
thick walled and verrucose at maturity, 1-2 celled, 5-7x35 µm, scar 2-2.5 µm, keratinolytic and cellulolytic.
38.C. vallenarense van Oorschot & Piontelli
Persoonia 12:487,1985.
Colonies restricted on YpSs at 25°C, white becoming sulphor yellow, conidia terminal, often developing
sympodially, rarely intercalary, tuberculate, 3.5-5.5x5-7
µm. Isolated from keratinous substrates. Conidia resembling those of Chrysosporium anamorph of Renispora flavissima.
39.C. siglerae Cano & Guarro Mycotaxon
51:75,1994.
10-15 mm on PYE in 21 days at 28°C, pale yellow,
conidia mostly lateral, 1 celled, smooth to slightly verrucose, 5-30x2-3.5 µm, 2 celled, 10-16x2-3 µm, intercalary
in series of two or more, 10-15x2-2.5 µm, keratinolytic.
40.C. farinicola (Burnside) Skou Friesia
11:70.1975.
27-45 mm in 14 days on honey agar, initially white
or becoming pale greenish yellow or pale brown, intercalary conidia abundant, smooth, thick walled, 3-12x5-12
µm, terminal and lateral smooth, thick walled, 1 celled,
6-15x4.5-9 µm, wide basal scar, 1.5-5 µm, not keratinolytic, osmophilic [22,23].
Species with uncertain position
41.C. parvum [or Emmonsia?]
42.C. crescens [or Emmonsia?]
Placed under Emmonsia Cif & Montemartini on
the basis of blastic conidia and thick walled chlamydospore- like celles [5,67].
43.C. racemosus Sharma, Bhattacharjee &
Bhadauria Indian Phytopath. 46:404,1993.
1-1.2 cms in 7 days at 28±1°C on PDA and
Czapecks dox agar, initially white, later changes to cream,
green, dark brown to blackish brown. Conidiophore branched with thick, numerous scars [hilum].
Acropleurogenous and pedicellate aleurospores aggregate
to form clusters at intercalry position of hyphae. The described species without camera leucida drawings or photographs does not seems to be Chrysosporium because
conidia do not have scars and are round. The cultures
were not available.
Species in press
44.C. undulatum Vidal, Ulfig & Guarro
45.C. fluviale Vidal, Ulfig & Guarro
46.C. submersum Vidal, Ulfig & Guarro
47.C. minutisporosum Vidal, Ulfig & Guarro
Genus Chrysosporium have following
teleomorphs also
Gymnoascus arxii, G. uncinatus, Nannizziopsis
spp., Amauroascopsis perforatus, Aphanoascus durus,
Aph. clathratus, Aph. fulvescens, Aph. hispanicus, Aph.
reticulisporus, Aph. saturnoideus, Aph. terreus, Aph.
verrucosus, Arthroderma multifidum, Arth. tuberculatum,
Arth. croccatum, Arth. silverae, Ctenomyces serratus,
Ajellomyces dermatitidis, Aj. capsulatus, Apinisia graminicola, Api. queenslandica, Neoxenophila foetida,
Renispora flavissima, Amauroascus albicans, Am. aureus,
Am. volatilis-patellis, Orromyces spiralis, Phaneochaeta
chrysosporoidea, Pseudarachmiotus orissi, Bettsia alvei,
Bettsia species, Cordyceps species.
Chrysosporium and its potential
Kushwaha RKS
69
Temperature
Mycelial growth of C. tropicum
There are many reports of the ability of
Chrysosporium to grow at sub zero and above 37°C.
C. keratinophilum, C. tropicum and C. queenslandicum
grows at 37°C. C. asperatum, C. pannorum and C. indicum grows in antarctic soil [68,69] and C. evolceanui in
alpine soil [70]. Garg et al [71] found 21-50 and 25-30°C
optimum for C. pannicola and C. keratinophilum. The
growth of C. tropicum was rapid at 27°C and 37°C showed minimum rate of growth [72]. Maximum germination
of conidia of C. tropicum took place at 32°C within 24
hours. Low temperature also supported conidial germination [72]. Slight growth of C. pannorum at -6°C [73] and
good growth at -5°C and at 5°C was reported [74-75].
Rapid growth of this fungus occurred at 15°C [76], maximum growth was at 25°C [74,75] and its rate of growth
reduced at 30-37°C [75,77,78].
Growth of C. tropicum was measured on 12 agar
media as follows: oat meal> YpSs> glucose asparagine>
potato dextrose> Czapecks dox> SAA> malt extract>
tryptone agar> Czapecks dox +yeast extract> peptone
dextrose> Sabouraud dextrose> Sabouraud dextrose+
yeast extract. Maximum sporulation was recorded on
Sabouraud dextrose agar supplemented with yeast extract
followed by tryptone agar [89]. This fungus grows in
varying concentrations of glucose while 6% glucose
favoured maximum growth. A study of carbon metabolism in C. tropicum was made at 5-15 days of incubation.
The rate of assimilation of glucose, sucrose, mannose,
maltose and lactose by C. tropicum was studied chromatographically [90]. Growth and sporulation of 15 species of
Chrysosporium including 6 strains of C. tropicum were
different on 7 media. Six strains could be categorised in
3 groups based on their growth characteristics [91].
Glucose supported maximum mycelial growth of
C. tropicum and fructose was assimilated more slowly
than mannose among the monosaccharides. Utilization of
sucrose, maltose and polysaccharide was average.
Mannose and starch showed an increase in growth rate up
to 10 days of incubation and then gradually decreased.
Mannose and maltose were utilized very rapidly by this
fungus and exhausted within 5 days. Glucose, fructose,
sucrose, lactose and starch were not completely utilized
[90]. C. tropicum synthesized galactose, glucose and fructose [90]. C. tropicum exhibited carbon heterotrophy, as
was reported in other soil saprophytes, and meets its
requirements from various sources.
pH requirements
A. uncinatum and A. curreyi are acidophilic and
C. tropicum, C. keratinophilum, and A. quadrifidum were
found to be alkalophilic [79]. C. pannorum grew well at
sea water salinity [77] but 20% sodium chloride inhibited
its growth when used in Czapecks medium [80]. A. curreyi survives in mud and sand dunes of coastal soils [81]
and C. tropicum and C. indicum in marine soils [82]. For
C. tropicum 7 pH was optimum but it grows at 3 and 7 pH
also [72]. Garg et al. [71] gave a range of soil pH in relation to the distribution of A. fulvescens, C. keratinophilum, C. pannorum, C. tropicum, C. asperatum,
C. evolceanui and C. serratus.
Moisture contents
Occurrence of A. curreyi was reported in nests
with 11.81% and 19.81% water content while C. keratinophilum was isolated from nests with higher moisture
content showing a hygrophilic nature [83]. The hygrophilic nature of A. fulvescens and C. keratinophilum was also
confirmed [84]. C. pannorum survived in habitats of little
or no biotic influence [68]. A high percentage of spore
germination in A. uncinatum and Ctenomyces serratus at
90-100 RH was recorded [85]. Minimum aW for growth
at 25°C on NaCl of C. pannorum, C. xerophilum and
C. fastidium was 0.92, 0.71 and 0.69 respectively. The
growth rate of C. fastidium in medium containing glycerol
was lower than with glucose and fructose. Pugh and
Evans [85] reported higher percentages of spore germination in A. uncinatum and C. serratus at 90-100 % RH.
Humus
Distribution of Chrysosporium was not affected by
humus [86] and this fungus along with C. indicum and
C. tropicum occurred in soil made up of disintegrated lava
with low organic matter [74]. Nigam and Kushwaha [23]
also reported C. carmichaelii, C. evolceanui, C. indicum,
C. keratinophilum, C. merdarium, C. pannicola, C. queenslandicum and C. tropicum in house dust, which is very
low in organic matter. Katiyar and Kushwaha [88] reported C. keratinophilum, C. tropicum and Chrysosporium
spp. with 100 % hair perforation ability from sand of
Mediterranean sea which was very poor in organic matter.
Nitrogen nutrition and metabolism
Analysis of filtrates of C. tropicum at 4 days revealed arginine, asparagine, aspartic acid, hydroxyl proline
and threonine, while at 12 days cystine, proline and serine
were also detected. Mycelial extract revealed the presence
of arginine, Y amino butyric acid, asparagine, cystine, histidine, hydroxy proline and serine.The asparagine utilization by C. tropicum was rapid. Suitability of ammonium
nitrate and some other nitrogen sources for the growth of
C. tropicum was studied by replacing asparagine. The rate
of utilization of asparagine and synthesis of cell bound
and cell free aminoacids was studied [92].
C. tropicum grew fairly well in nitrogen provided
in the form of nitrate but nitrate from ammonium source
supported completely less mycelial growth. Cystine supported maximum mycelial yield. Peptone was found to be
the best source for the fungus while tyrosine seems to be
poor in this respect. Alanine, arginine, aspartic acid, cystine, glycine, glutamic acid, histidine, leucine, methionine,
phenyl alanine, proline, serine, tyrosine, aspartic
acid+glutamic acid+arginine and alanine+ asparagine+
histidine+ phenyl alanine+ proline+ cystine and sodium
nitrate, ammonium sulphate and peptone were also all tested for the growth of C. tropicum.
Vitamin requirements
A mixture of biotin, cyanocobalmin, pyridoxin,
riboflabin was used with the omission of one vitamin each
time and the mycelial weight of C. tropicum was determined. It grew in the medium provided with all the five vitamins whereas it showed a sudden decrease when biotin
70
and cyanocobalmine were omitted individually from the
combination, pointing towards their deficiency [93].
Spore germination
Germ tubes of conidia of C. tropicum attained an
average length in plain agar in 24 hours while in SDA
mycelial clumps were developed in 24 hours. One percent
glucose and 0.01% peptone induced 100% germination
[94]. Maximum germination of conidia of C. tropicum
took place at 32 ºC within 24 hours. A low temperature of
12 ºC also supported conidial germination [94]. The keratinized and non-keratinized propagules of C. tropicum and
C. keratinophilum showed differences when germinated
on different substrates, but were similar in their ability to
tolerate temperature exposure by two methods [95].
Glucose was found to be a good carbon source for germination. Fructose, mannose, sucrose, maltose, lactose and
starch did not favour germination [94]. It was shown that
in C. tropicum with an increase up to 0.2% in the concentration of nitrogen content of sodium nitrate, the percentage of spore germination was also increased upto 60%
within 24 hours. Twelve per cent germination was recorded when 0.01% nitrogen from nitrate source was added
to the medium. Nitrogen given in the form of ammonium
sulphate exhibited an opposite effect. In this case the
maximum germination was recorded at 0.01% nitrogen. In
organic nitrogen, asparagine could induce germination up
to 52%. An equimolar mixture of asparagine, aspartic acid
and glutamic acid supported 58% spore germination of
C. tropicum [94].
Antibiotic response to growth and
spore germination
Dermostatin showed the highest inhibition of
C. tropicum at its lowest concentration of 200 µ g/ml
while aureofungin showed maximum inhibition at
1000 µ g/ml when supplemented with SDA. MICs of
aureofungin and dermostatin was 600 and 200 µ g/ml.
Maximum inhibition was caused by griseofulvin at
1000 µg/ml [96]. Complete inhibition of spore germination of C. tropicum was caused by aureofungin at a concentration of 500 µg/ml. Aureofungin was found to be
effective even at its lower dose as it also inhibits the
length of the germ tube. Dermostatin induced the swelling
of the spores before the emergence of the germ tube [96].
The culture filtrates of M. fulvum, M. gypseum and
T. mentagrophytes inhibited conidial germination of
C. tropicum [72]. Biomass of C. tropicum was reduced by
prednisolone at its lowest concentration [97].
Sensitivity to plant extracts, volatile
substances, soaps, detergents, oils and
biocides
Plant extracts of garlic, ginger, neem, ocimum,
onion, yellow oliander and a mixture of all these inhibited
growth of C. tropicum [72]. Mycostatic fumes of some
volatile substances such as formic acid, acetic acid, ethyl,
methyl, isopropile and butyl alcohols, diethyl ether and
chloroform were able to reduce the growth of C. tropicum
in liquid culture to more than half [72]. Biomass loss was
noticed of some keratinophilic fungi including C. tropicum by ethyl alcohol, isopropyle alcohol, chloroform, carbon tetra chloride, carbon disulphide, acetone and benzene
[98].
Opportunistic fungal strains have been found to be
implicated in mycotic diseases of man and animals.
Influence of some homeopathic drugs was studied to inhibit the hair invasion activity of A. terreus, C. keratinophilum and C. tropicum mechanically and ezymatically.
Mezerium, petroleum, ustilago and sepia caused 100%
inhibition of hair perforation and no protein could be released in the culture filtrates at their higher doses [99].
Similarily some hair dyes: black diamond, black rose,
mehndi and amla were used to inhibit hair perforation.
Complete inhibition of hair perforation was at 100 µg/ml
for black diamond and black rose, and the other two were
comletely inhibitory at 1000 µg/ml for these fungi. Five
Indian soaps: kesh nikhar, godrej shikakai, swastik shikakai, vipro shikakai were inhibitory for hair perforation at
1000 µ g/ml; shampoos: Optima, Organics, Pantene,
Sunsilk, detergents; Areil, Rin, Surf Excel, Wheel; and
agrochemicals: bavistin, thiram, neem, ocimum, urea,
rock phosphate, NPK and zinc sulphate were also completely inhibitory for hair perforation and protein release by
the above three fungi at their higher doses [100].
Six plant extracts of Lawsonia inermis, Eclipta
alba, Nyctanthus arbortristis, Datura stramonium and a
mixture of all the extracts were used for testing antifungal
activity of C. tropicum [101]. Essential oils of Mentha
arvensis, Trachyspermum ammi, Cymbopogan nardus and
Eucalyptus citriodora also caused more than 77% inhibition of C. tropicum [102]. Chrysosporium species were
inhibited by himax and tree burb [103]. Mycostatic effect
of tea, coffee, proteinex, coconut oil, linseed oil and vegetable oil was studied on C. tropicum. Coffee and tea
extracts were inhibitory for growth when these were used
with Sabouraud dextrose agar while proteinex showed
negligible inhibition, mustard and linseed oil were found
to be the most inhibitory [104]. C. pannorum was reported
to be resistant for higher concentration of organic mercurial preparation [105]. This fungus was also used in the
removal of phosphate from sewage [106]. C. keratinophilum was resistant to cadmium concentrations as high as
560 ppm [107].
Keratinolytic potential, enzymes and
secondary metabolites
C. pannicola, C. keratinophilum and C. tropicum
took as little as 5 days to colonize human hair [108].
C. carmichaelii, C. evolceanui and C. indicum were found
to be late colonizers of hair. A perforating group of
C. keratinophilum, C. pannicola, C. queenslandicum and
C. tropicum and a non - perforating group of C. carmichaelii, C. evolceanui and C. indicum were recognised.
C. indicum, C. keratinophilum and 2 Chrysosporium spp.
isolated from sand of a Mediterranean beach were able to
perforate human hair in 18 days and completely digested
it [88]. Five strains of C. tropicum caused 100 % hair perforation and 12 strains decolourised the hair, out of 12
C. indicum 5 caused 100% perforation and all 11 decolourised hair, of 10 C. queenslandicum 6 perforated hair and
all decolourised hair, of 5 C. keratinophilum 3 perforated
hair and all decolourised hair, of 3 C. pannicola all perforated hair and decolourised hair. All these strains when
grown on dermatophyte test medium develop zones ranging from 3-17 mm [109].
Fifteen strains of C. tropicum took 7-40 days for
colonization and perforation of human hair in soil [110].
In an another study C. carmichaelii, C. evolceanui,
C. farinicola, C. indicum, C. keratinophilum, C. lucknowense, C. pannicola, C. queenslandicum and C. tropicum
Chrysosporium and its potential
Kushwaha RKS
colonized human hair in vitro in 3-9 days [111]. Among
the eight species of Chrysosporium, C. keratinophilum
was able to perforate and degrade buffalo, cow, dog, goat,
horse and human hairs rapidly. Infected hair showed
undulation, lifting and disruption of cuticle, narrow and
broad perforating organs, projection of medulla and decolouration of hair as induced by this fungus [111]. English
[112] observed cuticle lifting, erosion of cortex and perforating organs in hair penetrated by C. keratinophilum.
Seven types of perforators developed by A. terreus,
C. keratinophilum and C. tropicum were also observed
[88,113-116]. The manner in which Chrysosporium species attacked hair was intermediate between dermatophytes and keratinophilic fungi [117].
C. keratinophilum, C. tropicum, C. indicum and
A. fulvescens were reported as keratin colonizers [118132]. Most active keratinolysis was shown by A. terrreus,
A. fulvescens, C. pannicola, C. queenslandicum, C. xerophilum, C. tropicum, C. keratinophilum, Chrysosporium
anamorph of A. cuniculi and Chrysosporium anamorph of
A. curreyi [133]. The penetration of hair in vitro by C. tropicum was similar to dermatophytes showing the ability to
infect [134]. The ability of A. fulvescens and A. verrucosus to penetrate hair, was found to be different from
A. keratinophilus [135]. Determination of this, and keratinolytic ability was demonstrated by a new method by inoculating the fungus directly on hair tied in a tube [136].
A. terreus, C. serratus, C. tropicum and C. keratinophilum isolated from museum soil were found to deteriorate feathers and produce 98, 44, 96 and 76 ku/ml
keratinase and release 680, 789, 830 and 650 µg/ml net
protein [137]. C. crassitunicatum and C. tropicum released 690 and 788 µg/ml protein and produced 88 and 33
ku/ml keratinase from hen feathers [138]. C. crassitunicatum, C. tropicum and C. indicum produced 130.2, 77.2
and 75 ku/ml keratinase when pig hairs were used [139]
and role of C. carmichaelii, C. evolceanui, C. indicum and
C. tropicum in keratin degradation [140] and characterization of extracellular proteolytic enzyme of C. tropicum
and its role in keratin degradation was also monitored
[141]. C. tropicum degraded buffalo horn, woman hair
and wool [142]. Amylase production by C. tropicum was
reported [143]. C. merdarium, C. keratinophilum, C. indicum, C. crassitunicatum, Chrysosporium anamorph of
Pectinotrichum llanense and Chrysosporium anamorph of
A. cunicuili were found to degrade chicken feathers [144].
Liquification of gelatin is a criterion for identification of the filamentous phase of Chrysosporium,
Blastomyces and Histoplasma [145]. C. indicum is able to
digest gelatin [146]. C. carmichaelii, C. evolceanui,
C. indicum, C. keratinophilum and C. merdarium, two
strains of C. queenslandicum, four strains of C. tropicum
also liquified gelatin [147]. C. tropicum, C. zonatum and
Chrysosporium anamorph of A. curreyi utilizes standard
lipids and fatty acids [cholesterol, palmitic and linolytic
acids] and evidence is available for the uptake and degradation of cholesterol by C. keratinophilum.
Nineteen enzymes were produced by 390 strains
of Chrysosporium [148]. C. tropicum- amylase, urease,
pectinase, keratinase, esterase lipase, leucine aryl amidase, cystine arylamidase, alpha galactosidase, alpha glucosidase, beta glucosidase, N acetyl glucosaminidase, alpha
mannosidase. C. merdarium- amylase, keratinase, urease,
esterase lipase, leucine arylamidase, alpha galactosidase,
beta galactosidase, alpha glucosidase, N acetyl glucosaminidase. C. indicum- amylase, cellulase, pectinase, keratinase, leucine arylamidase, cystine arylamidase, alpha
galactosidase, beta glucosidase, N acetyl-glucosaminidase, alpha mannosidase, C. keratinophilum- amylase, cellu-
71
lase, urease, pectinase, keratinase, esterase, esterase lipase, lipase, leucine arylamidase, chymotrypsin, alpha
galactosidase, beta glucuronidase, alpha glucosidase, beta
glucosidase, N acetyl glucosaminidase, alpha fucosidase.
C. queenslandicum- amylase, urease, pectinase, keratinase, esterase, lipase, leucine arylamidase, cystine aryla
midase, alpha galactosidase, alpha glucosidase, N acetyl
glucosaminidase. Chrysosporium anamorph of A. curreyiamylase, keratinase, esterase lipase, lipase, leucine arylamidase, cystine arylamidase, trypsin, chymotrypsin, alpha
galactosidase, beta glucuronidase, alpha glucosidase,
N acetyl glucosaminidase. C. carmichaelii- amylase,
urease, keratinase, esterase lipase, lipase, leucine arylamidase, cystine arylamidase, alpha galactosidase, alpha glucosidase, beta glucosidase, N acetyl glucosaminidase.
C. georgii- amylase, urease, pectinase, keratinase, esterase
lipase, lipase, leucine arylamidase, cystine arylamidase,
alpha galactosidase, beta glucosidase, N acetyl glucosaminidase.
Production of anthroquinines [questin and questinol] and asteric acid by C. merdarium was reported [149].
Secondary metabolites produced by keratinophilic fungi
were discussed [140]. Elastase was produced by C. evolceanui and C. indicum [150]. Phospholipids and acetone
soluble lipids were detected in cold mycelial extracts of
C. tropicum [151]. C. indicum produces L-Arginyl- D allothreonil- L-phenylalanine wich is antifungal [152].
C. pannorum produces cryscandin, antibacterial and anticandida and pinnorin-3-hydroxy-3-methylglutaryl co
enzyme, a reductase inhibitor [153,154].
Antagonistic potential
In vitro studies revealed that C. tropicum was inhibited by staling substances of C. evolceanui, C. indicum,
C. lucknowense, Aspergillus flavus, A. niger, Chaetomium
globosum, Cladospora species, three species of
Penicillium, T. vanbreuseghemii and M. gypseum. The
ability of C. tropicum to interact with 12 keratinophilic
and saprophytic fungi was evaluated in dual cultures. It
was overgrown by A. niger and C. globosum but other
fungi showed inhibition when in opposition to it. Eight
fungi were able to cause hyphal inhibition of C. tropicum
at a distance. Frequent curling, penetration, granulation,
lysis and chlamydospore formation in C. tropicum were
observed during hyphal interference. C. tropicum
penetrated the mycelium of A. niger [12]. Antagonistic
potential of C. tropicum against C. indicum, one
Penicillium species and M. gypseum is indicative of production of some inhibitory substances [12].
The growth of C. indicum was promoted by nine
keratinophilic fungi [155]. With C. tropicum it exhibited
maximum positive antagonistic potentiality while with
A. benhamiae - strain it showed least promotion in
growth. With both of these fungi C. indicum produced
intermingling colonies, the score being 1 and no zone of
inhibition. C. queenslandicum inhibited this to the maximum extent thus exhibiting a maximum negative antagonistic nature. The inhibiting species developed a zone of
inhibition exceeding 2 mm. This intermingled freely with
10 fungal partners while with 3 isolates it produced a zone
of inhibition less than 2 mm, the score being 2 as described for soil fungi [156]. C. queenslandicum showed the
greatest capacity to interact with as many as 14 fungi,
which it exhibited a promoting nature. Maximum antagonistic potential of C. queenslandicum to promote was
observed against Chrysosporium species with mutual inhibition and it was assigned a score of 2. This was more
inhibited when grown against Chrysosporium, an ana-
72
morph of R. vriesii, with the score being 3, Malbranchea
species could intermingle freely with it. Its higher index
of dominance showed its high cometitive nature.
The study of dual culture interaction showed that
out of 17 isolates interacted, two C. queenslandicum showed a similar antagonistic potential exhibiting a maximum antagonistic nature against other fungi. The overall
sequence of antagonistic potential can be summarized as
C. queenslandicum 264 and 265> Malbranchea sp.>
Chrysosporium sp. 234, Chrysosporium anamorph of
R. vriesii 208> Chrysosporium sp. 215, C. indicum,
C. indicum 201, 221, C. tropicum 263, 449> C. indicum
238> Chrysosporium sp. 442> Chrysosporium sp. 267>
A. benhamiae +> A. benhamiae -> A. ciferrii [155].
The antifungal potential of C. carmichaelii,
C. queenslandicum and six strains of C. tropicum was studied against Acrophialophora fusispora, Alternaria alternata, A. tenvissima, Aspergillus flavus, A. niger,
Aureobasidium sp., Botrytis sp., Cladosporium sp.,
Cunninghamella sp., Fusarium sp., Harposporium sp.,
Mucor sp., Penicillium citrinum, Rhizopus nigricans,
Torula sp., white sterile mycelium and three unidentified
fungi. One strain of C. tropicum allowed minimum number of fungi to grow. Staling substances of other
Chrysosporium spp. also caused inhibition of soil fungi.
Metabolites secreted by C. evolceanui, C. pannicola,
C. queenslandicum and one strain of C. tropicum showed
strong antifungal activity against A. niger [157].
Thirty Chrysosporium spp. caused inhibition of the
radial extension of M. gypseum, ranging from 20-100%.
Twenty one showed more than 50% inhibition of M. gypseum, 15 caused more than 50% inhibition of A. niger,
three strains completely inhibited A. niger, six caused
more than 50 % inhibition of P. citrinum while four completely inhibited this fungus. Neither of the C. evolceanui
were inhibitory. Out of eight strains of C. indicum three
inhibited M. gypseum. C. keratinophilum was less than
50% inhibitory. C. lobatum inhibited M. gypseum. One
strain of C. tropicum was 100% inhibitory for M. gypseum
and another for P. citrinum while other showed less than
50% inhibition of these fungi [155]. Interaction experiments on Chrysosporium anamorph of Arthroderma were
also carried out [158].
The volatiles emnated from three strains of C. tropicum were inhibitory for T. mentagrophytes. Inhibition of
T. rubrum was caused by volatiles of C. indicum, C. lobatum, and two strains of C. tropicum. The effect of fungal
staling substances of eight Chrysosporium spp. on soil
mycoflora was studied [157] and it was found that among
four strains of C. tropicum one allowed a minimum number of fungi to grow. Staling substances of other
Chrysosporium spp. also caused inhibition of soil fungi.
Metabolites secreted by C. evolceanui, C. pannicola,
C. queenslandicum showed strong antifungal activity
against A. niger [157]. The effect of staling products of
keratinophilic and non-keratinophilic fungi on the growth
and spore germination of C. tropicum was studied [159].
The inhibition /promotion in dual cultures depends on
many factors such as staling products of interacting colonies, pH change, nutrient media depletion or alteration of
nutritional ingredients besides hyphal interference.
Colony interaction in a number of cases is represented by
mutual inhibition in growth of both fungal partners. The
relative difference reveals the measure of susceptibility
and antagonistic ability of the fungus. The development
and subsequent formation of chlamydospores, lysis, coiling, deformation, granulation and swelling in dual cultures in most cases are evidence of their successful
competition [14].
Combined effect of Chrysosporium
with other keratinophilic fungi on hair
and feather decomposition
Combination of C. keratinophilum with C. queenslandicum, C. tropicum and M. gypseum enhanced feather
decomposition. The synergistic action of C. keratinophilum and M. gypseum was most effective as it caused highest protein release in the course of feather decomposition
in vitro. The course of wool degradation by C. keratinophilum acting singly and in combination with C. carmichaelii, C. tropicum and M. gypseum was studied by
measuring protein released in the culture medium and
weight loss of wool up to four weeks. The combined
effect of these fungi on the continual breakdown of wool
by C. keratinophilum and the effect of the addition of
C. keratinophilum on the continual degradation of wool
by C. carmichaelii, C. tropicum and M. gypseum were
also studied. The synergistic action of C. keratinophilum
and M. gypseum on wool was found to be more effective.
The biodegrading potential of C. keratinophilum can be
made more effective by M. gypseum and C. carmichaelii,
if the latter fungi join halfway through keratinolysis.
C. keratinophilum did not act as a follower fungus in wool
degradation [167]. A series of experiments were performed to see if keratinolytic C. carmichaelii and C. tropicum could be effective on the amount of wool
decomposed by C. keratinophilum. It would be expected
to occur because some of these might be utilising protein
released as it breaks down. In doing so, it should relieve
competitive inhibition of keratinase action and also relieve
repression of keratinase synthesis [167].
C. indicum, C. keratinophilum, C. pannicola,
C. queenslandicum and C. tropicum were also used for
their capacity to degrade horn, hair and wool [142].
C. europae decomposed feather and released
457.33 µ g/ml protein along with another 47
Chrysosporium strains tested. NaCl, KCl and Ca Cl2 inhibited protein release from feathers when C. keratinophilum was used. This fungus was also used for feather
decomposition during solid state fermentation [169].
Fungal colonization of hair in contact
with soil
C. tropicum appeared in 6 days on fresh feathers in
peptone and water amended soil while it took more time
in the presence of glucose to colonize defatted and sterilized feathers. In the presence of water 16-20 days were
taken to colonize 100% fresh and defatted feathers. More
than 25 days were required for achieving 100% colonization in the presence of glucose. Sterilized human hair was
not completely colonized in peptone and glucose amended
soil up to 25 days. Cow, goat and horse hairs were colonized completely in 20 days while fresh horse hair required
more than 25 days in glucose amended soil [111].
Five strains of A. keratinophilus perforated hair in
soil and caused 48-54% hair perforation in 30 days [100].
A. terreus perforated 63-100% hair. C. articulatum and
C. carmichaelii colonized hair in 12 and 18 days but could
not perforate them. Six C. indicum colonized hair in 4-15
days and 5 strains perforated hair. Five strains each of
C. keratinophilum, C. pannicola and C. tropicum colonized 100% hair in 3-5 days. C. queenslandicum, C. zonatum and C. xerophilum also colonized hair through soil
but later did not perforate hair.
Chrysosporium and its potential
Kushwaha RKS
Decomposition of hair and feather
in soil
The hair from humans and animals and feather
from birds which come to the soil either as dropped off or
dead are affected by microbial decomposition. In the past
few decades some studies on the decomposition of keratin
in submerged cultures appeared. Biodegradation of keratin
by using Chrysosporium and other related fungi in submerged culture is reviewed [160] and scattered reports are
available in literature [137,138,142,161-168,170-172].
The process of keratin decomposition has also been found
to be very fast in soil and it plays a very important role in
energy transformation and nutrient cycling in soil.
Decomposition of keratin in soil has received very little
attention [167,174,175]. Several species of
Chrysosporium were found to degrade hair through soil
by a new method [166]. Combinations of C. indicum and
C. keratinophilum, C. pannicola and C. keratinophilum,
C. queenslandicum and C. keratinophilum, C. tropicum
and C. keratinophilum, M. gypseum and C. keratinophilum were employed for finding out weight loss of hair and
feather in soil
The midway addition of the C. keratinophilum in
C. queenslandicum experimental set showed 100% weight
loss of feathers in 3 weeks. It is evident from overall
observations made during experiments that, in general, the
fungi acting in combination are found to be more effective
in keratin decomposition than the individual action of
various fungi. The midway addition of C. keratinophilum
was the most effective in C. tropicum amended soil while
midway additon of C. queenslandicum was more successful when initially inoculated with C. keratinophilum.
Feather and wool degradation by C. keratinophilum was studied in natural garden and sterilized garden
soil, where it was presumed that C. keratinophilum utilises keratin without any competition from microorganisms
for up to 3 weeks, and later a slight decline was noted.
This showed that C. keratinophilum can efficiently cause
keratin breakdown while competing with other fungi in
soil [167]. The effect of C. tropicum on soil which had
received decomposed wool products was studied during
seed germination and seedling growth of Brassica campestris Linn, Zea mays Linn and Phaseolus Roxb. and it
was concluded that this fungus could be used for the slow
release of nitrogen fertilizer in soil [176].
Pathogenicity
Chrysosporium sp. was cultured from a tissue
biopsy of the nasal mucosa which was found in brain,
lungs and left kidney as well as nasal and sinus regions
[177]. Chrysosporium sp. was isolated from 2 biopsy specimens of a 24 - year - old man [178] and three other
male patients [179].
Pathogenecity of Chrysosporium spp. and C. parvum var crescens was also confirmed [180-182].
Chrysosporium infection in a bone marrow transplant
73
recipient was noted as Chrysosporium which caused an
invasive infection in an 18 - year - old woman where
infection began as a facial swelling and which extended
into the central nervous system [183]. Studies were
carried out on the epidemiological, immunological, biochemical and physiological properties of Chrysosporium
sp. and C. keratinophilum along with some dermatophytes
[184].
The pathogenic role of C. keratinophilum, C. asperatum, C. georgii, C. tropicum, C. pannorum,
Chrysosporium state of A. curreyi, Chrysosporium state of
A. multifidum, Chrysosporium state of A. tuberculatum is
uncertain but their ability to remain viable for several
weeks in skin and peritonial tissue indicates that they
could become pathogenic in certain circumstances
[181,182]. Antigenic activity within the genus
Chrysosporium
was
demonstrated
[185].
A. keratinophilus, A. fulvescens, A. reticulisporus and
A. verrucosus produced nodular lesions when inoculated
intraperitonially [186]. The isolation of C. merdarium
from nail, C. pseudomerdarium from lung of rodent,
C. carmichaelii from human skin and sputum, C. queenslandicum from snake, Chrysosporium anamorph of
Gymnoascus demonbreunii from human, Chrysosporium
anamorph of Pectinotrichum llanense and C. inops from
human skin, C. sulfureum from bones, Chrysosporium
anamorph of Rollandina vriesii from skin and lung of
lizard, C. lobatum from scrappings of human and skin
crest and C. pannicola from dog [5] are all also indicative
of their potential for pathogenecity.
Future Prospects
It is felt that there has been no attempt to determine the geographical distribution of Chrysosporium. Large
areas of the world have yet to be sampled for its teleomorphic connections and germplasm collection.
Molecular characterization of Chrysosporium species, its
teleomorphs and related genera may be able to provide
help in solving diagnostic problems but this criterion
should be cosidered as the second step because visualization of characteristics has always had the upper hand.
More information on the pathogenic and saprophytic survival, spread and nature of the life cycle of
Chrysosporium is needed. An approach in using combinations of wild as well as genetically engineered strains of a
single species or different species of Chrysosporium for
enhancing biodegrading potential merits study. Detailed
ecological and physiological studies followed by suitable
selection and development of specific strains may lead to
a commercially viable use of fast - growing non - pathogenic strains of this fungus.
Most of the work cited here is financed by
Department of Science and Technology and
University Grants Commission, New Delhi, India.
74
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