MICROBIAL CULTURES preservation

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Microbial Cultures: Maintenance, Preservation and Registration
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2. REVIEW
National Agriculturally Important Microbial Culture Collection
in the Global Context of Microbial Culture Collection Centres
Sushil K. Sharma1 • Sandeep Saini1 • Ankita Verma1 • Pawan K. Sharma1 •
Rup Lal2 • Manish Roy1 • Udai B. Singh1 • Anil K. Saxena1 • Arun K. Sharma1
Received: 26 September 2016 / Revised: 3 May 2017 / Accepted: 18 May 2017
The National Academy of Sciences, India 2017
Abstract Understanding the metabolic and evolutionary
patterns of microorganisms has played a pivotal role in the
development of agriculture, industry and health sectors.
Therefore, for the ex situ conservation of the microbial
diversity, microbial culture collections also known as Bio-
banks or Microbial Resource Centres remain the most
important scientific infrastructure. This review describes the
history and evolution of microbial culture collections and the
growth of the global community of collections through the
activities of the World Federation for Culture Collections
(WFCC). In addition, it highlights the roles of culture col-
lections in assisting research and development including the
role of an International Depository Authority recognized
under Budapest Treaty. Furthermore, the status of microbial
culture collections available in India with emphasis on col-
lection of agriculturally important microbes has been inves-
tigated. National Agriculturally Important Microbial Culture
Collection is a designated national repository established at
the Indian Council of Agricultural Research–National Bureau
of Agriculturally Important Microorganisms, Maunath,
Bhanjan, Uttar Pradesh, India in 2004 and is an affiliate
member of WFCC (WDCM-1060) which currently encom-
passes over 6000 well characterized strains of bacteria,
cyanobacteria, fungi etc. The deposited microorganisms are
being used as bioinoculants, biopesticides and for manage-
ment of soil fertility, biotic and abiotic stresses in crops for
sustainable production.
Keywords Culture collection
International Depository Authority Microbial diversity
Extreme environment
Introduction
Microorganisms are a natural resource that exist in all
habitats even in hostile climatic conditions where virtually
all life cannot exist. Although these microbes are very
small, they constitute the highest percentage of living
organisms on earth [1, 2]. So far, only a small fraction of
this vast diversity has been explored and in order to
facilitate the expansion of the current microbial diversity
available for research, their long-term preservation is
needed. To date the majority of microorganisms cannot be
grown in culture and many function as part of integrated
microbial communities unable to grow without others. It is
an essential role of microbial collections to make these
organisms, as well as those isolated available to the
research community. Once isolated from their natural
habitat, it is generally required that the microorganisms be
cultured for conservation and utilization. These microor-
ganisms play pivotal roles in the functioning of ecosystems
to maintain a sustainable biosphere. They have been used
as biological control agents, biocatalysts and produce a
wide variety of natural products (e.g., drugs, enzymes,
Significance statement The paper traces history of microbial culture
collections and highlights role of the National Agriculturally
Important Microbial Culture Collection (NAIMCC) in conservation
of microorganisms for their utilization in agriculture and allied
sectors. It also underpins the problemsrelating to sharing of microbial
resources among different countries.
Sushil K. Sharma
sks_micro@rediffmail.com;
http://www.nbaim.org.in
1
National Agriculturally Important Microbial Culture
Collection (NAIMCC), ICAR- National Bureau of
Agriculturally Important Microorganisms,
Kushmaur, Maunath Bhanjan, Uttar Pradesh 275103, India
2
Molecular Biology Laboratory, Department of Zoology,
University of Delhi, New Delhi 110007, India
123
Proc. Natl. Acad. Sci., India, Sect. B Biol. Sci.
DOI 10.1007/s40011-017-0882-8

3. metabolites, etc.) that have agricultural, pharmaceutical
and industrial applications [3]. They are decomposers,
converting nutrients from organic wastes and dead organ-
isms into molecules that can be reused within ecosystems
and are recognized as vital components of the world’s
biodiversity. Besides nutrient cycling, these microorgan-
isms are also involved in establishment of beneficial
mutual relationships and in the production of atmospheric
oxygen [4–6]. This versatility in function has brought
microorganisms to the fore, turning them into the key
players in the dawning of the biotechnological era of the
20th century. It is important to emphasize that microor-
ganisms are an essential part of the natural elemental cycle
and a source of novel products for application in agricul-
ture, medicine as well as nutrition and therefore their
preservation is as important as it is for any other biological
species [7–9]. In the past, despite such vast importance, not
much attention has been paid to the conservation of
microorganisms in many developing countries including
India. Microorganisms have been isolated from natural
habitats since the time of Pasteur, yet only a small fraction
of these has been preserved throughout globe. Large
numbers of microorganisms have been left unpreserved and
cultures lost due to unavailability of suitable conditions for
preservation as well as lack of knowledge regarding the
process of conservation [10, 11]. In addition, loss of
microbial cultures in the past can also be attributed to
unavailability of adequate permanent and reliable micro-
bial culture collections. Microbiologists isolate, select,
investigate, classify, improve and genetically modify dif-
ferent natural strains isolated from different habitats. Due
to limitations imposed in the isolation methods as well as in
culture conditions, some strains once isolated may not be
re-isolated. Therefore, in this perspective efforts must be
made to deposit these isolated novel microorganisms to the
culture collections in order to maintain their integrity for
screening, characterization, genetic improvement and pro-
duction of useful products for the scientific community in
the future [10, 12, 13]. In this review, the authors attempt to
highlight the progressive status, history and evolution of
microbial culture collections in the world, presenting the
NAIMCC of India as a key example.
History and Evolution of Culture Collections
Bacterial and Fungal Culture Collections
The first public service bacterial culture collection was
established by Prof. Frantisek Kral at the German University
of Prague, Czech Republic. He worked as a technician in
1890 at the institute of Hygiene; German University of
Prague [14]. Prof. Soyka chose him as curator of the bacterial
culture collection and he was later appointed as Associate
Professor of Bacteriology. Based on his experience in han-
dling microorganisms, he brought out the first catalogue of
microbial strains from the bacterial culture collection in 1900
(Fig. 1). After Kral’s death in 1911, the culture collection
was then transferred to the State Serum Institute, Vienna. The
former director of Serum Institute, Prof. Ernst Pribram joined
Loyola University in Chicago, Illinois in 1927 and brought
part of the Kral bacterial collection with him. During World
War II, the culture collection left in Vienna was destroyed.
Later on, other culture collections were also established after
Kral’s collection [15]. Public service collections of the period
were Mycotheque de I’ Universite Catholique de Louvain
(MUCL) set up in 1894 at Louvain –la Neuve, Belgium and
the collection of the Centraal Bureauvoor Schimmelcultures
(CBS) in Utrecht, the Netherlands in 1906, both focussing on
fungi. Afterwards, more microbial culture collections were
established. Some collections were of general purpose, while
others were specialized culture collections that dealt with
specific kinds of microorganisms. The American Type Cul-
ture Collection (ATCC) was originally established in
Washington in 1925 and is now located in Manassas, Vir-
ginia, USA. The CABI collection, formerly the Common-
wealth Mycological Institute collection, was founded as a
UK National Resource in 1947 by consolidating its scientists’
collections maintained from its origins in 1920; this collec-
tion provided a resource to support both fungal taxonomy
and agriculture globally [16]. The DSMZ-German Collection
of Microorganisms and Cell Cultures was established in the
year 1969 as the National Centre for Culture Collection in
Germany and subsequently recognized by International
Depository Authority (IDA) in 1981. Prof. Julio R. Vil-
lanueva initiated the establishment of the Spanish Type
Culture Collection (CECT, Coleccion Espanola de Cultivos
Tipo) in Valencia, Spain in 1960. At the outset, the CECT
was located in the Biological Research Centre (CIB) of
Spanish Research Council (CSIC), Madrid, Spain. After-
wards it was shifted to Department of Microbiology,
University of Salamanca in 1967 by Prof. Villanueva. Later
in 1974, the CECT was transferred to Department of
Microbiology, University of Basque Country, Lejona, Viz-
caya, Spain and Prof. Fedrico Uruburu became director of
the culture collection. The first microbial strain catalogue
was made available in 1976. The CECT became a member of
WFCC in 1977 and finally in 1980, the CECT was housed in
the research building of the University of Valencia. In 1981,
European curators, in the fourth meeting of WFCC decided
to establish the European Culture Collections’ Organization
(ECCO). The CECT joined ECCO in 1983 and was recog-
nized as an IDA in 1992 under the Budapest Treaty (Fig. 1)
[15, 17]. There have been several publications that have
partially addressed the history of collections looking at their
development [18–20]. There have been a number of
S. K. Sharma et al.
123

4. publications that address the collections from their functions
perspective [21, 22] and also trace the history of single col-
lection [23]. The significance of the history of collections is
in collection development enabling one to learn from this to
help accelerate progress and to be able to focus on providing
high quality resources with appropriate associated data to
meet future needs. For example, recent papers look at the
development and funding of the next generation culture
collections the microbial domain Biological Resource Cen-
tres (mBRC) [24–28]. Lists of collections and their back-
ground can be found at several websites, a few of them given
below.
http://www.mycology.net/Content/MNetContent.cfm?MID
=135
http://www.wfcc.info/index.php/collections/display/
http://mycology.cornell.edu/
Algal Culture Collections
In the beginning of the 20th century, due to increasing sci-
entific interest in the study of algae, some specialized culture
collections were established. The first Algal Culture Col-
lection was established at Charles University in Prague
where Horejsi (in 1910) successfully isolated symbiotic
cyanobacteria from Cycas revoluta (Fig. 1). Uhlir
(1892–1915) recovered cyanobacteria from the lichen Col-
lema. Cultures were maintained in vitro using a mineral
nutrient solution. After the death of Uhlir in 1916, Prat
followed a similar path and started isolating and cultivating
Oscillatoria. Prat continued his studies on algal physiology
in the Laboratory of Plant Physiology at the Czech part of
Charles University in Prague. Unfortunately, the majority of
cultures were lost due to lack of financial support [29].
During the 1920’s a Centre of Cultivation of Algae was
developed at the German part of Charles University of
Fig. 1 Culture collection timeline. The timeline highlights the major events in the history of culture collection till the establishment of National
Agriculturally Important Microbial Culture Collection (NAIMCC)
National Agriculturally Important Microbial Culture Collection in the Global Context of…
123

5. Prague by Ernst George Pringsheim who made significant
contributions to modern phycology [30, 31]. Pringsheim
published the first catalogue comprising information of
about 49 strains held in the culture collection. On account of
political turmoil, Prof. Pringsheim left Charles University of
Prague and went to England in 1938 with some strains and
established a Culture Collection at Cambridge University,
UK, now named the Culture Collection of Algae and Pro-
tozoa (CCAP). Internationally, CCAP is a member of the
WFCC and ECCO. After his time as a refugee scientist,
Pringsheim came back to Germany in 1953 and established
a Culture Collection of Algae at Gottingen University
(SAG) which is now an affiliate member of WFCC (Fig. 2)
[32]. This culture collection comprised primarily of micro-
scopic algae and cyanobacteria. The Pringsheim cultures
were also used for establishing the Culture Collection of
Algae at the University of Texas at Austin (UTEX) in the
USA (1953) by R.C. Starr, who was Pringsheim’s collabo-
rator at Cambridge University [33]. After the end of World
War II, Prat again went back to the culture collection of
Prague University with his collaborators. Later, Fott added
the collection of green algae and heterokonts to the culture
collection of Prague University. The major part of the cul-
ture collection was transferred to Trebon in 1979 and has
now been established as Culture Collection of Algae at the
Laboratory of Algology (CCALA), Czech Republic. The
CCALA is a member of both Federation of Czechoslovak
Collections as well as WFCC.
The Belgian Co-ordinated Collection of Microorgan-
isms (BCCM) is the only culture collection specialized in
conservation of diatoms. The BCCM was originally
established in 2000 by Dr. V.A. Chepurnov at Laboratory
for Protistology and Aquatic Ecology (PAE), Gent
University, Belgium which is internationally renowned for
its research on taxonomy, evolution, ecology and life
cycles of diatom. As with the bacterial and fungal collec-
tions, a lot can be learned from the already established
collections equally true for the microalgae collections
where organisms can be more difficult to preserve. Bring-
ing the collection communities together can accelerate the
discovery of solutions to the many challenges faced.
Culture Collections: The Global Scenario
The WDCM website on 31 December 2016 listed
713 culture collections from 74 countries or regions hold-
ing 1051,947 bacteria, 788,397 fungi and totalling more
Algae at the
Culture Collection of
University of Texas at
Austin (UTEX),
established in 1953 by
R.C.Starr, U.S.A.
Algae at Gottingen
University (SAG),
established in 1953 by
Prof. E.G. Pringsheim
Germany.
Algae at Laboratory
of
Algology(CCALA),T
rebon,established in
1979 by Fott.
of Algae
Protozoa (CCAP),
UK established in
1947 by Prof.
E.G.Pringsheim
Algae, Germany
established in 1920 by
Prof. E.G. Pringsheim)
Culture Collection of Culture Collection of
Culture Collection
Culture Collection of
Fig. 2 Origin of four major algal culture collections from culture collection of algae established in 1920 by Prof. E.G. Pringsheim, Germany
S. K. Sharma et al.
123

6. than 2.5 million microorganisms. Among the culture col-
lections, 232 from Europe, 250 from Asia, 181 from
America, 41 from Oceania and 11 from Africa are regis-
tered with WDCM (http://www.wfcc.info/ccinfo/statistics/
). The use of collections can be assessed by the Analyzer of
Bio-resource Citations (ABC) which finds collection strain
numbers in publications; currently138,408 papers provide
reference to 74,340 strains, belonging to 131 of the WDCM
registered culture collections in 50 countries, these have
been published in 50,307 journals since January, 1953 (
http://abc.wfcc.info/). The Global Catalogue of Microor-
ganisms (GCM) is a user-friendly tool by which culture
collections manage, disseminate and share the information
pertaining to their holdings [34]. The WFCC Executive
Board has recently updated the WFCC Guidelines for the
establishment and operation of Collection of Cultures of
Microorganisms. The dissemination of the WFCC guide-
lines at local level reinforces the awareness and improves
the communication among members locally and abroad.
The WFCC organizes the International Congress of
Culture Collections (ICCC) every 4 years and the 13th
meeting of ICCC was organized in 2013 in Beijing, China
and the 14th ICCC is scheduled for 30 October to 5th
November 2017 in Antalya, Turkey (http://www.wfcc.
info/news/culture_collection). The WFCC publishes
newsletters providing time to time updates of the registered
culture collections and also accelerates the information
related to new advances in preservation of microorganisms.
It also compiles the information about the different confer-
ences, workshops and meetings held in different countries
on topics such as the recent advancement in culture col-
lections and current status of the conservation strategies of
microbial diversity and their biotechnological applications.
The WFCC supports its member collections to respond to
new challenges including those of the regulatory environ-
ment, for example compliance with the Nagoya Protocol
which came into force on 12th October, 2014. The protocol
contains rules for access to genetic resources and the fair and
equitable sharing of the benefits arising out of their utiliza-
tion (http://www.wfcc.info/pdf/wfcc-nl-december-2014-sc.
pdf). India is implementing this protocol through the
National Biodiversity Authority (NBA), Ministry of Envi-
ronment, Forest and Climate Change (MoEF CC), Govt.
of India for exchange of germplasm under the umbrella of
Biodiversity Act 2002 and Rules 2014.
Origin of WFCC and Bioinformatics
for Systematics
In 1962 during a conference held under the aegis of
International Union of Microbiological Societies (IUMS), a
section was set up specifically for culture collections which
finally came into force in 1963. A committee was set up to
regulate this section with Chairman Prof. Skerman (Aus-
tralia) and his colleagues. The constitution for the section
was further revised by the committee members in 1966 in
Paris. In a subsequent meeting in Tokyo in 1968, it was
proposed to convert the section into the World Federation
for Culture Collections (WFCC) with an integration of new
laws. Finally, in 1970 during Xth International Congress
for Microbiology held in Mexico City, the section voted to
dissolve and was reconstituted as the WFCC which has
been active since then [35]. WFCC is a multidisciplinary
international federation, which coordinates the activities of
culture collections worldwide. With the exhaustive diver-
sity of microorganisms, it is essential to maintain an
inventory in order to demonstrate the heterogeneity of the
cultures and the data associated available for research. The
WFCC supports the World Data Centre of Microorganisms
(WDCM) in the maintenance of an online international
database that compiles data on the culture collection cen-
tre, management, services provided and most recent
research supported using informatics. The WDCM pro-
vides a single interface to access the comprehensive
microbial resource information by the scientific and
industrial communities [36, 37]. Currently, the WDCM
coordinates information from over 700 collections around
the world. With the ever-growing field of Next Generation
Sequencing (NGS) technologies, genome data for
microorganisms is becoming more readily available. There
is a need to coordinate access to the genomic blueprint
along with other features of the microorganisms. As of
today, WDCM offers three bioinformatics tools for easy
access to culture collections related data namely, InfoBIO,
G-InforBIO, and O-InforBIO (http://www.wfcc.info/links/
). Each of these tools is dynamic in nature and provides
user-friendly interfaces for microbial data analysis and
comparative genomics. While InfoBio stays focussed on
data basing the information from culture collections,
G-InforBIO, and O-InforBIO provides platforms for
accessing genome annotations and designing customized
databases, respectively.
Acquiring the Status of Depository Institutions
as Under Budapest Treaty
Any country which respects intellectual property rights has
its own national policy and provision for patenting of an
invention. In 1973, the British Government proposed that
the World Intellectual Property Organization (WIPO),
Geneva, Switzerland should initiate ‘‘single deposit’’ ful-
filling the requirement of depositing cultures in other
countries where patent applications are filed. The objective
was to simplify procedures for seeking patent protection
National Agriculturally Important Microbial Culture Collection in the Global Context of…
123

7. for inventions involving microbes in more than one country
and to evolve a uniform system and guidelines. In April
1977, a conference was organized in Budapest, Hungary,
on the recommendation of experts and a treaty was adopted
which is called the ‘‘Budapest Treaty on the International
Recognition of the Deposit of Microorganisms for the
Purposes of Patent Procedure.’’ The Budapest Treaty came
into force in 1980 [3]. Under the Budapest Treaty, certain
culture collections were recognized as ‘‘International
Depository Authorities (IDAs).’’ There are 44 International
Depository Authorities (IDAs) in 24 countries worldwide
and majority of them are in Europe http://www.wipo.int/
export/sites/www/treaties/en/registration/budapest/pdf/ida.
pdf. As defined in the treaty, a deposit of a microorganism
associated with a patent can be made in any one IDA
which is recognized and accepted by all the contracting
states for patent procedures. Each individual IDA can
decide its own scope for the type of microorganisms it
would like to accept. For example, an IDA like the
Deutsche Sammlung von Mikroorganismen und Zell
Kulturen (DSMZ, Germany) accepts bacteria, fungi, bac-
teriophages, plasmids, plant viruses, plant cell cultures,
human and animal cell cultures and murine embryos. The
Czech Collection of Microorganisms (CCM), Czech
Republic accepts bacteria, filamentous fungi, yeast-like
microorganisms and yeasts whereas American Type Cul-
ture Collection (ATCC, USA) accepts bacteria, viruses,
yeasts, fungi and protozoa (Source http://www.wipo.int/
treaties/en/registration/budapest/summary_budapest.html).
There are two IDAs in India, the Microbial Type Culture
Collections (MTCC, India) accepts actinomycetes, bacte-
ria, fungi, yeasts and plasmids and the Microbial Culture
Collection (MCC), National Centre for Cell Science
(NCCS) which accepts bacteria, fungi, yeasts and plasmids
in a host and/or as isolated DNA preparations belonging to
Hazards Group 1 and 2 as per classification of the Indian
Authority; genetically manipulated microorganisms and
isolated DNA are accepted if they can be processed in
BSL-1 or BSL-2 facility or conform to Group 1 or 2
organisms. WIPO made available the 2015 statistics for
microorganism deposits in August 2016 at http://www.
wipo.int/budapest/en/. There have been a total of almost
97,000 deposits since the Treaty came into force (almost
5000 in 2015 alone) with over 31,000 held in the ATCC
and almost 8000 in the DSMZ. Growth has been noted in
China, with the China General Microbiological Culture
Collection Center (CGMCC) now holding almost 12,000
microorganisms associated with patents. If India can
facilitate exchange of microorganisms internationally,
there is potential to grow in this field too, currently the
MTCC holds 850 strains with the MCC holding 60.
Housing Agriculturally Important
Microorganisms
Agriculturally important microorganisms are key to the
sustaining of agriculture globally. The agricultural research
aims to improve crop production, nutritional security and
economy utilizing beneficial microorganisms. Plants, ani-
mals, fishes, insects and microorganisms are valuable
genetic resources for future development of human civi-
lization. It is of paramount importance to conserve and
sustainably use agro-biodiversity which is declining rapidly
due to changing climatic situations. Agriculturally impor-
tant culture collections in this context play an important
role in providing authentic microbial cultures to research-
ers working in different domains of biology. Some of the
culture collections which hold microorganisms of agricul-
tural importance are listed in Table 1. CABI is an organi-
sation dedicated to agriculture and holds 28,000 strains
including 6000 plant pathogens and many other soil borne
organisms of relevance to soil health in particular and to
agriculture in general. Agricultural Culture Collection of
China (ACCC) was established in 1980 and is one of the
Culture Collection under China Committee for Culture
Collections of Microorganisms (CCCM). It now contains
more than 4100 strains including bacteria, actinomycetes
and fungi. Culture Collection of Beijing Agriculture
University holds around 5060 Rhizobium strains which are
kept both in freeze dried form and in mineral oil. Korean
Agricultural Culture Collection (KACC) with holdings of
4900 was established in 1995 to conserve microbial
diversity for utilization in agriculture. Indian Type Culture
Collection (ITCC) was established in 1936 whereas both
National Rhizobial Germplasm Collection and Centre for
Conservation and Utilization of Blue Green Algae
(CCUBGA) were established in 1986 in the premises of
Indian Institute of Agricultural Research (IARI), New
Delhi. Centre for Mycorrhiza Culture Collection (CMCC)
under The Energy Resource Institute, New Delhi is a
Mycorrhizal Bioresource Centre which aims at conserva-
tion of mycorrhizal biodiversity by means of collection,
propagation, characterization and maintenance of cultures
under in situ conditions. The Centre for Mycorrhizal Cul-
ture Collection (CMCC) was established in 1993 which
seeks support from the Department of Biotechnology,
Govt. of India. Since then, the bank has a glorious col-
lection of over 600 different isolates of which 257 are
ectomycorrhizal fungi (EMF) and over 350 are Arbuscu-
lar Mycorrhizal Fungi (AMF) isolates. International Cul-
ture Collection of Vesicular-Arbuscular Mycorrhizal Fungi
(INVAM) is located at West Virginia University Research
Collection. INVAM is the largest culture collection which
S. K. Sharma et al.
123

8. contains 99% unique strains. INVAM currently contains
1112 accessioned strains comprised of 112 species sub-
mitted from 49 countries.
Microbial Culture Collections: Indian Scenario
India is recognized as one of the hotspots of megabiodi-
versity encompassing enormous endemic microbial
diversity. Extensive bio-prospecting projects in the past
funded by the Department of Biotechnology revealed the
immense microbial diversity which could potentially be
exploited for medicinal, industrial and agricultural pur-
poses. Currently, India houses 30 microbial resource
centres (MRCs) that are registered with the WDCM. Such
a large number of culture repositories are in coherence
with the microbiologically diverse nation that India is.
With more than 194,174 strains currently deposited at
various MRCs, India holds the third highest number of
strains behind the USA and Japan. With the advent of
microbial and genomics era, an increasing number of
MRCs began to be recognized as IDAs, first among them
being the Microbial Type Culture Collection (MTCC),
Chandigarh, India, recognized as IDA in 2002. MTCC
houses more than 9000 strains in total comprising bacteria,
fungi, yeast and actinomycetes. Another national reposi-
tory is the Microbial Culture Collection (MCC), Pune,
holding more than 150,000 microbial strains [3]. The
MCC additionally provides various other services includ-
ing identification, biochemical characterization, 16S rRNA
gene sequencing, phylogenetic analysis, MALDI-TOF,
FAME and DNA–DNA hybridization. A fungal conser-
vation facility, National Fungal Culture Collection of India
(NFCCI), Pune, India was established by the Department
of Science and Technology in 2008. The NFCCI is an
exclusive repository holding over 2950 strains of different
groups of fungi. Some other WFCC registered culture
collections include National Collection of Industrial
Microorganisms (NCIM), Pune with 2950 strains, Indian
Type Culture Collection (ITCC), New Delhi with 3820
strains, National Collection of Dairy Cultures (ICAR-
NDRI), Karnal with 460 strains and Bank A Bug (BAB),
Gandhi Nagar with 3373 strains.
Sharing the Microbial Biodiversity of India
The primary aim of MRCs is to conserve and manage
microorganisms, cell lines, genetic and other biological
materials and to make them accessible to researchers for
harnessing their potential for the benefit of society. The
Convention on Biological Diversity (CBD;www.cbd.int)
places sovereign rights over genetic resources with the
Table
1
Some
agriculturally
important
microbial
culture
collections
in
the
world
Source
modified
from:
http://www.wfcc.info/ccinfo/collection/by_id/97,
http://www.mycology.net/Content/
MNetContent.cfm?MID=135,
http://www.wfcc.info/index.php/collections/display/,
http://mycology.cornell.edu/
S.
no.
Name
of
culture
collection
Country
Number
of
microbial
strains
1
Agriculture
Culture
Collection
of
China
(ACCC)
China
4100
bacteria,
fungi
and
yeast
strains
2
Centre
for
Agriculture
and
Biosciences
International
(CABI)
United
Kingdom
26,000
fungal
and
2000
bacterial
strains
3
Culture
Collection
Beijing
Agricultural
University
(CCBAU)
China
5060
Rhizobium
strains
4
Collection
for
Extremophile
Microorganisms
and
Type
Cultures
(EMTC)
Russian
Federation
1595
bacterial
strains
5
Russian
Collection
of
Agricultural
Microorganisms
(RCAM)
Russian
Federation
5104
strains
6
Korean
Agricultural
Culture
Collection
(KACC)
Korea
17,612
bacteria
and
fungal
strains
7
National
Collection
of
Agricultural
and
Industrial
Microorganism
Hungary
3312
bacteria,
fungi
and
yeast
strains
8
International
Collection
of
Microorganisms
from
Plants
New
Zealand
18,675
bacteria
and
fungi
strains
9
United
States
Department
of
Agriculture
(USDA)
America
1310
bacteria
and
fungi
strains
10
National
Agriculturally
Important
Microbial
Culture
Collection
(NAIMCC)
India
6000
bacteria,
fungi
including
cultures
repatriated
from
CABI
cyanobacteria
strains
11
Agricultural
Research
Service
Culture
Collection
(NRRL)
USA
96,198
bacteria,
fungi
and
yeast
strains
12
Centre
for
Mycorrhiza
Culture
Collection
(CMCC)
India
More
than
600
Mycorrhizal
strains
13
International
Collection
of
Vesicular
Arbuscular
Mycorrhizal
Fungi
(INVAM)
United
States
1112
strains
14
Indian
Type
Culture
Collection
(ITCC)
India
More
than
4000
fungal
strains
National Agriculturally Important Microbial Culture Collection in the Global Context of…
123

9. country of origin. India takes the position that material
transfer agreements (MTAs) of all MRCs bind the recipient
to use the material only for research purposes. For com-
mercial utilization, prior permission from the competent
authority is mandatory along with a benefit sharing
arrangement with the country from where the resource
originates. Keeping this in mind, the CBD (Rio de Janeiro,
1992) adopted the Nagoya Protocol on 29th October, 2010
in Nagoya, Japan. The Nagoya protocol was implemented
from 12th October, 2014 with access to genetic resources
and fair and equitable sharing of benefits arising from their
utilization as the major objective. As of the 31st December
2016, the Access and Benefit Sharing Clearing House
indicates that there are 86 parties to the Nagoya protocol,
79 countries have ratified the protocol, and India is a sig-
natory (https://absch.cbd.int/). In the eleventh meeting of
the conference of parties, held in Hyderabad, India rein-
forced that ‘‘nature protects if it is protected’’ (https://www.
cbd.int/doc/meetings/cop/cop-11/official/cop-11-35-en.pdf).
The competent body to monitor implementation of Nagoya
protocol in India is the National Biodiversity Authority
(NBA), Ministry of Environment, Forest and Climate
Change, Govt. of India. Recent regulations of the NBA,
dated 21st November, 2014 placed restrictions on the
export of microbial resources outside the country without
prior permission of the NBA even if the microbial
resources are to be utilized for research purposes only. This
proved to be a great hurdle as far as microbiological
research is concerned. NBA regulations have sparked
nationwide discussions and in response, the NBA has eased
the deposition of novel taxa to foreign culture collections in
the 37th Authority Meeting by replacing ‘‘prior permission
of the NBA’’ with ‘‘prior intimation to the NBA through
Form-C’’ for novel taxa deposition. However, foreign
researchers still need approval from the NBA before
ordering strains of Indian origin from international repos-
itories, even for research purposes. Due to this regulatory
enforcement by NBA, researchers and culture collections
of India are now facing two serious problems (1) type
strains deposited in Indian culture collections are not
recognised as valid deposit for publication purpose since
they are not available to researchers abroad and (2) foreign
culture collections refuse to accept microbial cultures from
Indian researchers because of clause of seeking ‘‘prior
approval’’ from NBA. To help researchers performing
taxonomical characterization, the culture collections in
India can take the initiative to share cultures worldwide by
securing permission from NBA instead of the researchers
having to seek that permission themselves. In order to
facilitate compliance with the requirements of the NBA,
there must be a streamlined process and it would be in best
interests of research that culture collections centralize this
process with NBA.
Although it is our onerous responsibility to protect and
preserve the national biodiversity, it is also essential to plan
implementation of national laws and regulations in a way
that does not hamper ongoing research. The standard MTA
of MRCs places the buyer of biological material under
obligation to use the biological material only for research
purposes. The MTA makes it mandatory for the recipient to
act according to the benefit sharing requirements of the
Nagoya protocol. The NBA regulations have placed strict
restrictions on sharing of biological material even for
research purposes even when MRCs through their MTA
have ensured all recipients are required to act according to
the Nagoya protocol. This is doubly ensured as recipients in
signatory countries will be required in their country to have
in place appropriate benefit sharing agreements and have
legitimately accessed the genetic resources they use. The
whole idea of MRCs takes a backseat in India when we
cannot share our biodiversity with the world. It is impera-
tive to say that the country of origin has benefit rights over
biological material, but restricting biodiversity is not always
equivalent to conserving biodiversity. Also, it is almost
impossible to restrict the transfer of microscopic entities
such as bacteria. A person’s microbiome, consisting of
billions of bacteria is exported outside the country every
time the person decides to travel out of the country. It is thus
crucial to debate whether ubiquitous entities such as the
microbes can be regulated in terms of export and import and
whether such stringent regulations are actually needed to
preserve the biodiversity? MRCs would lose their purpose
eventually if researchers cannot even share their knowledge
and materials freely with the world and discovery of
microorganisms from our own country would be difficult
because type cultures are mostly available in MRCs located
in other parts of globe. Therefore, NBA should develop
mechanisms that facilitate India’s collaboration with
researchers abroad and should address the issues associated
with research harmonization with the rest of the world. The
Nagoya Protocol principles are being implemented globally
but processes put in place by the nations to implement these
principles can be very different (see in view of some facts
highlighted by Overmann and Scholz [38]. It is very
important that communities and nations introduce mecha-
nisms to harmonise approaches to facilitate compliance in a
way that does not impede science and the benefits it can
provide. The Microbial Resources Research Infrastructure
(MIRRI) has produced the MIRRI Access and Benefit
Sharing Manual which addresses best practice for microbial
domain Biological Resource Centres, this manual can be
downloaded from the MIRRI website (http://www.mirri.
org/fileadmin/mirri/media/Dokumente/generalDocs/MIRRI_
ABS_Manual_web.pdf). Further best practices from other
communities can be found on the ABS Clearing House
website https://absch.cbd.int/.
S. K. Sharma et al.
123

10. Genesis and Establishment of NAIMCC
Evolution
ICAR-NBAIM was established by the Department of
Agricultural Research and Education (DARE) and ICAR,
Ministry of Agriculture, Government of India under the IX
Five Year Plan in 2001. The initial foundation of ICAR-
NBAIM was at ICAR-National Bureau of Plant Genetic
Resources (ICAR-NBPGR), New Delhi in 2001. The entire
Bureau was moved and established as a separate unit in the
empty/unoccupied building of National Institute of Sugar-
cane and Sugar Technology (NISST) at Kushmaur, Mau-
nath Bhanjan, Uttar Pradesh on 1st June, 2004. The DARE
and ICAR jointly laid the foundation of ICAR-NBAIM
with underlying objectives to explore, collect, maintain and
conserve microbial wealth of the country and to ease the
availability of agriculturally and industrially important
microorganisms for researchers, farmers, industry and
others stakeholders. A landmark development of ICAR-
NBAIM was the simultaneous establishment of NAIMCC
in the year 2004. CABI helped the collection in its estab-
lishment by providing protocols, exchanging scientists and
supplying many reference strains of Indian origin. The
storage capacity of NAIMCC currently is nearly 10,000
agriculturally important microorganisms (AIMs) with fur-
ther expansion in rapid progress. The NAIMCC was reg-
istered as an affiliate member of WFCC in 2014
(www.wfcc.info/ccinfo/collection/by-id/1060) and is strin-
gently built on the guidelines and principles as laid down
by the WFCC/OECD. The ICAR-NBAIM has been rec-
ognized as a Designated Repository (DR) by the NBA,
MoEFCC, Govt. of India (www.nbaindia.org) for man-
agement of agriculturally important microbial genetic
resources of the country.
Current Status
NAIMCC holds 6322 accessioned microbial strains com-
prising of 3801 fungal strains, followed by 2293 bacterial
strains and 228 cyanobacterial accessions. Successive
increase in microbial holdings with every progressing year
is represented in (Table 2). In a broader perspective, 50
bacterial genera, 33 fungal genera and 18 cyanobacterial
genera constitute the total microbial holdings in NAIMCC
(Table 3). Amongst the microbial cultures deposited in
NAIMCC; Bacillus, Pseudomonas, Streptomyces, Lacto-
bacillus and Paenibacillus represents the prominent bac-
terial genera, fungal genera chiefly comprise of Fusarium,
Trichoderma, Pleurotus, Aspergillus, Alternaria, Agaricus,
Colletotrichum, Chaetomium and predominant cyanobac-
terial genera include Nostoc, Anabaena and Calothrix.
Table 2 Year-wise increase of culture holdings in NAIMCC since
2009
Years No. of culture holdings
2009 2495
2010 2753
2011 3842
2012 4448
2013 4690
2014 5243
2015 6017
2016 6157
2017 6322
Table 3 Bacterial, actinomycetes, fungal and cyanobacterial genera present in NAIMCC
S.
no.
Microbial groups Name of genera
1 Bacteria and
actinomycetes
Actinoplanes, Acinetobacter, Azotobacter, Aeromonas, Arthrobacter, Alcaligenes, Agrobacterium, Azospirillum,
Bacillus, Brevibacillus, Bradyrhizobium, Brevibacterium, Burkholderia, Citribacter, Citrococcus,
Cellulomonas, Edwardsiella, Enterococcus, Enterobacter, Escherichia, Exiguobacterium, Frankia, Geobacillus,
Haloarcula, Halobacillus, Halomonas, Klebsiella, Lysinibacillus, Lactococcus, Lactobacillus, Mesorhizobium,
Microbacterium, Micrococcus, Methylobacterium, Ochrobactrum, Pseudomonas, Paenibacillus, Pantoea,
Ralstonia, Sinorhizobium, Rhizobium, Sphingobacterium, Stenotrophomonas, Sphingomonas, Staphylococcus,
Streptococcus, Serratia, Streptomyces, Vigibacillus and Xanthomonas.
2 Fungi Agaricus, Alternaria, Acremonium, Aspergillus, Beauveria, Curvularia, Colletotrichum, Cochliobolus,
Cladosporium, Chrysosporium, Chaetomium, Fusarium, Glomerella, Ganoderma, Hypsizygus, Mucor,
Microascus, Macrophomina, Metarhizium, Pleurotus, Paecilomyces, Penicillium, Phoma, Rhizopus,
Saccharomyces, Sclerotium, Schizophylium, Thielavia, Trichoderma and Verticillium
3 Cyanobacteria Aulosira, Anabaena, Aulosira, Cylindrospermum, Chrococcus, Calothrix, Hapalosiphon, Lyngbya, Microchaete,
Nostoc, Oscillatoria, Plectonema, Pscillatoria, Phormidium, Scytonema, Spirulina, Tolypothrix and
Westiellopsis
National Agriculturally Important Microbial Culture Collection in the Global Context of…
123

11. Table 4 Some representative microorganisms possessing agriculturally and industrially important traits
S. No Traits Name of microorganisms Accession number
1. Nitrogen fixation Bradyrhizobium sp. IC 7114
Azotobacter chroococcum1A
NAIMCC-B-00239
NAIMCC-B-00045
2 Plant growth promotion
HCN, ammonia and ligninase producer Stenotrophomonas maltophilia AMAAS-137
Enterobacter cancerogenus AMAAS-198
NAIMCC-B-00946
NAIMCC-B-00951
IAA and siderophore producer Agrobacterium tumefaciens VA9S9
Acinetobacter sp. VA2S2
NAIMCC-B-01084
NAIMCC-B-01086
Protease, cellulase, amylase, siderophore producer Bacillus pumilus NFB-3
Pseudomonas tolaasii NNB-4
Bacillus amyloliquefaciens GA-8
NAIMCC-B-01219
NAIMCC-B-01221
NAIMCC-B-01223
Phosphate solubilizer Streptomyces griseolus ARHSPO17
Penicillium albicans CABI-233074
Trichoderma harzianum CABI-288012
Pseudomonas taiwanensis MPF-2
NAIMCC-B-00914
NAIMCC-F-01427
NAIMCC-F-01731
NAIMCC-B-01337
Zinc solubilizer Burkholderia arboris NKD-11
Bacillus tequilensis JUKD5
NAIMCC-B-01383
NAIMCC-B-01392
Ligninase producer Bacillus subtilis subsp. subtilis AMAAS157 NAIMCC-B-00948
Phytate mineralizer and zinc solubilizer Bacillus aryabhattai MDSR14
Bacillus aryabhattai MDSR07
NAIMCC-B-01442
NAIMCC-B-01440
Bioagent Trichoderma longibrachiatum 31PP NAIMCC-F-02938
3. Biotic stress management
Biocontrol of soil/seed borne pathogen Acremonium strictum CABI-230422
Bacillus atrophaeus SJ13
Bacillus pumilus P72
Dactylaria sp.CABI-327745
Streptomyces avidinii VH-32
NAIMCC-F-00052
NAIMCC-B-01222
NAIMCC-B-00849
NAIMCC-F-00716
NAIMCC-B-00603
Entomopathogen Bacillus thuringiensis sub sp israelensis
Aspergillus fumigatus CABI-016195
Beauveria bassiana CABI-189552
Metarhizium anisopliae CABI-15223
Bacillus thuringiensis sub sp. aizawai NCIM 5116
NAIMCC-B-00124
NAIMCC-F-00262
NAIMCC-F-00389
NAIMCC-F-01291
NAIMCC-B-00121
4. Abiotic Stress Management
Cold tolerant Bacillus pumilus NBA1
Bacillus malacitensis NBO1
Paenibacillus xylanexedens NBJ5
Brevibacterium frigoritolerans NBC7
Janthinobacterium sp. L110
NAIMCC-B-00771
NAIMCC-B-00779
NAIMCC-B-00774
NAIMCC-B-00785
NAIMCC-B-01122
Thermotolerant Pseudomonas putida AKMP7
Paenibacillus macerans NBH-7
Aquitalea magnusonii NBRAJG84
Acinetobacter baylyi NBRAJG74
Acinetobacter baumannii NBRAJ73
NAIMCC-B-00922
NAIMCC-B-00710
NAIMCC-B-00030
NAIMCC-B-00013
NAIMCC-B-00003
Salt tolerant Pseudomonas pseudoalcaligenes CIBA-T3
Alteromonas macleodii CIBA-AG-2
Halomonas sp. NBSL-19
Halobacillus dabanensis CIBA-CST4
Marinobacter alkaliphilus NBSL-7
NAIMCC-B-00512
NAIMCC-B-00513
NAIMCC-B-00616
NAIMCC-B-00578
NAIMCC-B-00610
Osmotolerant Enterobacter asburiae TRM
Enterobacter asburiae P17
NAIMCC-B-01340
NAIMCC-B-01341
S. K. Sharma et al.
123

12. NAIMCC’s accessioned bacterial species belong mainly to
the phylum Firmicutes followed by c-Proteobacteria,
Actinobacteria, a-Proteobacteria. The bacterial culture
diversity alone has been deposited from 25 different states
of India. Supplementing this further, fungal and
cyanobacterial cultures deposited here represents the
widespread collection from 23 to 19 different Indian states,
respectively. Accessioned fungal genera mainly belong to
classes Deuteromycetes, Basidiomycetes, and
Ascomycetes. Most of the cyanobacterial holdings at
NAIMCC belong to the orders Chroococcales, Nostocales,
Stigonematales and Oscillatoriales with maximum mem-
bers belonging to order Nostocales and Oscillatoriales. Of
the 228 accessioned cyanobacterial genera Nostoc, Ana-
baena, Calothrix and Oscillatoria are greatest in number. It
is pertinent to mention that amidst the total fungal cultures
deposited, 1427 culture vials are ones that have been
repatriated from CABI. Recently, NAIMCC has also
Table 4 continued
S. No Traits Name of microorganisms Accession number
Industrially Important Lactobacillus fermentum B2-8
Lactococcus lactis sub sp. lactis BHS1
Lactobacillus plantarum ADF10
Limnothrix sp. 01
Anabaena doliolum 28
Plectonema radiosum 25
Plectonema notatum 34
Lyngbya sp. 12
NAIMCC-B-01196
NAIMCC-B-01319
NAIMCC-B-01051
NAIMCC-C-00107
NAIMCC-C-00002
NAIMCC-C-00026
NAIMCC-C-00120
NAIMCC-C-00111
Table 5 Preservation methods of microbial cultures in NAIMCC
Methods of preservation
Short-term methods Long-term methods Other methods
1. Sub-culturing
• Live pure microbial cultures stored at 4 C in
culture medium
• Culture incubated aseptically in sterile agar slants
with periodic transfer
• Many bacteria and fungi can be kept for 3-4 months
• Use of low concentration medium prolongs time-
interval of sub-culturing
• Cultures lose viability faster than frozen stocks
1. Freeze drying (lyophilization)
• Bacteria and spore-forming fungi can be preserved
effectively by lyophilization for long-term storage
• Lyoprotectant such as skim milk (20%) and sucrose
(5–10%) are used as freeze drying solutions
• Method includes freezing, primary drying and
secondary drying
1. Storage of cultures in
sterile distilled water
2. Vitrification of obligate
parasites and fruiting
body of mushroom fungi
3. L-drying for
microorganisms
4. Storage of cyanobacterial
culture under active
cultivation
5. Silica based storage of
cyanobacteria
6. Substrate based
lyophilization of
mushroom fungi
7. Encapsulation of microbes
for cryopreservation
8. Paper disc method for
storage of yeast
9. Filter paper method for
storage of spore-forming
fungi
2. Mineral oil storage
• Method of choice for preservation of fungal
cultures
• Oil reduces oxygen level for cultures and thereby
decrease growth of cultures
• General survival rate of culture is 2–3 years
• Appropriate for fungal plant pathogens as it
maintains the pathogenicity of fungi.
2. Cryopreservation
• Preservation of microorganisms in liquid nitrogen at
-196 C or in gas phase above liquid nitrogen at
-150 C along with croprotectants
• Method provides long-term survivability and stability to
bacteria and fungi
• The non-sporulating basidiomycetes and mycorrhizal
cultures can be cryopreserved as they are susceptible
to lyophilization
3. Deep freezing
• Method of storage of bacteria and fungi
• Cultures preserved using a cryoprotectant (glycerol
or DMSO) to reduce damage from ice crystal
formation
• Bacteria and fungi preserved in 16–20% (v/v) and
10% glycerol, respectively followed by storage at
20/80 C in a deep freezer
• Cultures can be preserved in glycerol stocks for
1–2 years
National Agriculturally Important Microbial Culture Collection in the Global Context of…
123

13. received 475 cultures of mushroom from ICAR-Directorate
of Mushroom Research (ICAR-DMR), Solan and approx-
imately 2000 cultures are yet to be received. Some of the
cultures have also been received from ATCC (American
Type Culture Collection), USA, NCL, Pune, CCUBGA,
New Delhi and MTCC, India. At present in India,
NAIMCC represents the only culture collection with an
exclusive focus on preservation of microorganisms pos-
sessing agriculturally and industrially important traits such
as phosphate solubilisation, production of siderophore,
cellulase, ligninase and antibiotic production and mitiga-
tion of biotic and abiotic stresses such as drought, cold,
high temperature, salt etc. (Tables 4, 5).
Services
NAIMCC accepts microbial cultures under general and safe
deposit. The microbial germplasm registration facility is also
available at NAIMCC for registering elite microbial germ-
plasm which is distinct, unique, and stable and has potentially
proven attributes of academic, scientific, agricultural and
industrial importance. The Microbial Germplasm Registra-
tion Committee (MGRC) is the advisory committee which
approve the registration of the microbial germplasm. The
details of submission of microbes for accession number and
supply of cultures to users are available at Microbial Genetic
Resource Portal (www.mgrportal.org.in). Recently, ICAR-
NBAIM has attained the designation of a nodal body by
Central Insecticides Board and Registration Committee (CIB
and RC) for storage of registered and to be registered
biopesticides in NAIMCC (cibrc.nic.in). The diagrammatic
representationofthe activitiesofNAIMCCisshown inFig. 3.
Linkages
NAIMCC has countrywide linkages with Indian Type
Culture Collection (ITCC, New Delhi); National Dairy
Research Institute (ICAR-NDRI, Karnal), Microbial Type
Culture Collection (MTCC, Chandigarh), Microbial Cul-
ture Collection (MCC, Pune), National Collection of
Industrial Microorganisms (NCIM, Pune), National Fungal
Culture Collection of India (NFCCI, Pune), Veterinary
Type Culture Collection,(VTCC, Hisar), Cyanobacterial
Culture Collection, Imphal (IBSD) and Centre for Con-
servation and Utilization of Blue Green Algae (CCUBGA),
New Delhi [39].
Conclusion
India has made significant progress in microbial resource
management after the Convention of Biodiversity (CBD)
came into existence in 1992 and it was realized that
microorganisms represent a ‘‘black box’’ that has potential
solutions to various problems pertaining to food, health,
environment and poverty in both developed and developing
countries. It is clear that these resources are not only
valuable to India but could play a major role in the global
context. Before the CBD, many developing countries
including India did not give much attention to the conser-
vation of microbial resources but now repositories of India
are developing strategies and methodologies to conserve
the vast microbial wealth of the country. MRCs play a
crucial role in the characterization, conservation and sus-
tainable use of microorganisms in agriculture and allied
sectors. Researchers require reliable cultures for their
research; therefore, there is a strong desire for the long
term preservation of microbial cultures for present and
future use. In this context, NAIMCC plays a crucial role in
maintenance, conservation and utilization of microbial
diversities of agricultural importance in India. A current
constraint in microbial research in India is the restriction
imposed by the National Biodiversity Authority (NBA) on
exchange of cultures of Indian origin with the international
scientific community. Failure to provide type strains/ref-
erence strains to investigators outside India for research
purposes puts Indian Culture Collections at a disadvantage
as currently, culture collections abroad neither recognise
type strains deposited in Indian Culture Collections as a
‘valid deposit’ because they are not available to researchers
abroad nor are they accepting cultures of Indian origin
because of the clause ‘prior approval’ from NBA. Indian
microbial repositories, namely NAIMCC, MTCC, MCC
and others are working with the NBA to address the need to
ease the regulations for research purposes under the Bio-
diversity Act 2002 of India to enable Indian scientists and
NAIMCC
Safe
deposit
Germplasm
registration
Deposit of
biopesticide
under
CIBRC
Services for
identiication
and
characteriza
tion
Supply of
cultures in
active
form
Receipt of
request for
cultures
General
deposit
Deposit of
cultures
Fig. 3 Overview of activities of NAIMCC
S. K. Sharma et al.
123

14. research institutions to participate in international research
initiatives. These discussions have succeeded to an extent
that NBA has developed Form C for depositing the
microorganisms in non-Indian repository for claim of novel
taxa but still a lot more needs to be addressed.
Acknowledgements The authors are grateful to Dr. D. J. Bagyaraj,
CNBRCD, Bengaluru for critical reading and suggestions given to
improve the quality of the manuscript. SKS is grateful to Dr. D. L.
N. Rao, ICAR-IISS, Bhopal for his encouragement to consolidate
information on importance of culture collections that deal with agri-
culture including algal culture collections developed by Prof. E. G.
Pringsheim. The authors express their gratitude to Dr. Manjit Singh
and Dr. R. C. Upadhyay, ICAR-Directorate of Mushroom Research,
Solan, H.P, India for their help in transfer of mushroom cultures to
NAIMCC. We thankfully acknowledge the contribution and input of
Dr. David Smith, Centre for Agriculture and Biosciences International
(CABI), Egham, Surrey TW20 9TY, United Kingdom, in the final
draft of MS. They are thankful to Mr. Alok Upadhyay, Senior
Technician for his time to time help in providing information relating
to NAIMCC.
Compliance with ethical standards
Conflict of interest The authors declare that they have no conflict of
interest.
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