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Fungi: Their role in deterioration of cultural heritage
Department of Biotechnology, University of Natural Resources and Applied Life Sciences Vienna, Muthgasse 11, 1190 Vienna, Austria
a r t i c l e i n f o
Received 28 October 2009
Received in revised form
17 March 2010
Accepted 18 March 2010
a b s t r a c t
Fungi play a considerable role for the deterioration of cultural heritage. Due to their enor-
mous enzymatic activity and their ability to grow at low awvalues fungi are able to inhabit
and to decay paintings, textiles, paper, parchment, leather, oil, casein, glue and other
materials used for historical art objects. The weathering of stone monuments is signifi-
cantly increased by epi- and endolitic fungi. In museums and their storage rooms, climate
control, regular cleaning and microbiological monitoring are essential in order to prevent
fungal contamination. Education and close collaboration of mycologists and restorers are
needed to develop object specific methods for the conservation and treatment of contam-
ª 2010 The British Mycological Society. Published by Elsevier Ltd. All rights reserved.
Fungi – including yeasts, moulds, mushrooms and toadstools
– have been playing a considerable role for human culture and
the evolution of human society for thousands of years. Yeasts
wereused forbeer and bakeryproductsby the Egyptians,Kelts
and Teutons – albeit not being aware of microbiological
processes. Mushrooms probably served as food for hunters
and gatherers since the beginning of humankind and toad-
stools as Amanita muscaria and species of Psilocybe were used
as hallucinogens for cultic rites form Sibiria to South America.
The psychedelic effect of LSD 45 derived from Claviceps pupur-
eum influenced authors – Ernest Ju ¨nger, Aldous Huxley – and
somehow even catalysed the so-called ‘‘cultural revolution’’
in the 1960s. A real revolution of invaluable benefit was the
detection of the first antibiotic by Flemming in 1930 which
laid the basis for a completely new life-saving therapeutic
approach. The history of fungi in medicine and their revolu-
tionary impact in the cure of infectious diseases was recently
highlighted by Wainwright (2008).
In contrast to their numerous beneficial effects, fungi also
have their ‘‘dark side’’: mycotoxins, pathogenicity, allergens,
food spoilage and biodeterioration of materials. Biodeteriora-
tion of houses by mould was already mentioned in the bible
as white, red or green ‘‘leprosy’’ or ‘‘fretting’’ on brick, clay
and wood (Old Testament, Third Book of Moses, chapter 14,
verses 33–57). Today, fungal contamination is an increasing
problem not only in houses and working places. Objects of
art in museums and their depots are seriously threatened by
fungal contamination. The prevention of mould growth in
museums as well as the development of appropriate treat-
ment measures for contaminated objects is a challenge for
* Corresponding author.
E-mail address: firstname.lastname@example.org
1749-4613/$ – see front matter ª 2010 The British Mycological Society. Published by Elsevier Ltd. All rights reserved.
journal homepage: www.elsevier.com/locate/fbr
fungal biology reviews 24 (2010) 47–55
Author's personal copy
restorers, museum curators and architects. This has implica-
tions for the techniques of cleaning and conservation of
objects but also consequences for the occupational safety
and health of restorers and other museum personnel. In
outdoor environments fungi and lichen are the most impor-
tant agents of biodeterioration of historic monuments and
sculptures made of stone, mortar and plaster.
2. Fungi in the museum environment
Contamination of pieces of art presented in exhibition rooms
or stored in depots and their spoilage by fungi is not excep-
tional but rather frequent in old and newly built museums
(Allsopp et al., 2004; Nitte ´rus, 2000a; Capitelli et al., 2009;
Manoharachary et al., 1997; Mesquita et al., 2009; Pangallo
et al., 2007; Koestler et al., 2003). It is well known to mycologists
that fungi are able to inhabit, to alter and to degrade all types
of organic and inorganic materials (Fig. 1A–F). However, most
conservators andmuseumcuratorsarenotawareofthis enor-
mous deteriorative potential.
Historically, pieces of art were made of all types of organic
materials and these materials are again used for an authentic
restoration or conservation of the objects in recent times:
Paint was made of mineral pigments bound with organic
binders such as egg yolk, casein, linseed-, poppy seed-, hemp-
seed oil, Chinese wood oil or different resins. Linen canvas
clamped on wooden frames serves as painting ground and
was often primed with rabbit skin glue before painting. Gold
Fig. 1 – (A, B) Part of a wood ceiling with 12th century paintings. The paint layer was deteriorated by Aspergillus sp. (C) Pastel
painting with fungal contamination due to packing in plastic foil. (D) Mould on imperial Austrian horse trappings made of
textiles. (E) Historical frames with gold leaf strongly contaminated by fungi. (F) Dense lawn of Trichoderma sp. on historical
48 K. Sterflinger
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leaf on precious wooden or stucco frames was applied using
organic glues, linseed or turpentine oil. Historic glues were
based on cellulose or rabbit skin. Sculptures and other art
objects carry de ´cors made of textiles, leather, straw, clay,
natural hair or feathers. The most precious documents of
humankind are books and scrolls made of paper, papyrus
and parchment. Because of the tremendous diversity of exo-
enzymes produced by fungi – cellulases, glucanases, laccases,
phenolases, keratinases, mono-oxygenases and many more –
and their remarkable ability to grow at low awvalues the pres-
ervation of museum objects is inevitably connected with
prevention of mould, monitoring of mould and treatment of
mould on contaminated objects.
ings and other materials in museum is given in Table 1. The
data are based on more than 20 studies carried out in Austrian
museums since 2000 and on data collected from different case
mycetes; hemiascomycetes – yeasts – are rarely isolated from
art objects. Teleomorphs are rarely found and the only teleo-
morph genera that frequently occur are Chaetomium – mostly
on paper, wood and feathers – and Eurotium – in environments
with low awvalues. Occurrence of basidiomycetes is restricted
to wood degradation in churches or other protected historical
monuments. Zygomycetes are frequently isolated from pieces
of art but in most cases they can be regarded as transients not
being really established on the objects.
Fungal growth on objects of cultural heritage often causes
a serious aesthetical spoiling due to colony formation and
fungal pigments (Sterflinger et al., 1999). Moreover, fungi
degrade materials and thus affect objects substantially: the
enzymatic degradation of organic binders causes reduction
or even loss of paint layers (Fig. 1A–C). Fungi penetrate cracks
and migrate underneath paint layers thus causing detach-
ment. In paper conservation fungi are a special problem due
to their ability to excrete cellulases (Fig. 1F). Lignin degrading
fungi are rarely observed in indoor environments, but consid-
erable damage can be caused by the cellulose degraders Ser-
pula lacrymans or Conophora puteana in churches and other
objects of cultural heritage if wooden altars or the roof struc-
tures are attacked. Also originals and museum reconstruc-
tions of historical buildings are considerably damaged by
S. lacrymans (Bech-Andersen and Elborne, 2004).
The development of fungi in museums is to a large extent
determined by the indoor climate, the amount of available
selves – and also by the cleaning intervals in the museum. The
indoor climate as indicated by temperature, relative humidity
growth. It is also closely related to the buildings physical prop-
erties, especially the thermal insulation and the tendency to
generate condensate from warm indoor air on cold walls of
the building envelope (Camuffo, 1998). Depending on the
climate in the museum or storage rooms the fungal diversity
is restricted to few xerophilic and xerotolerant species such
as Eurotium sp., Aspergillus sp. or Wallemia sp.. Only in storage
rooms where the humidity is raised to more than 70 % for
a period of several weeks or month is a high fungal diversity
able to establish. The climate ranges allowing fungal spores
to germinate and that restrict the growth of the fungal myce-
lium are shown in the isopleth systems by Sedlbauer and
Krus (2003). The authors also show that hygroscopic materials
support the growth of fungi at low relative humidity and that
the water demand depends on the biodegradability of the
substrate. Theobjectsinfluencethedevelopment ofthefungal
community by their chemical composition and biodegrad-
ability for species with different exo-enzymes.
In museums the range of 55 % RH is generally regarded as
the border line for fungal growth and thus climate control is
adjustedbelow this value. In fact, fungi that are able to survive
at a relative humidity of 55% are rare and restricted to extreme
environments such as hot and cold deserts. So why do meso-
philic hyphomycetes frequently occur in museums? All
museums in the world measure temperature and humidity
in storage and exhibition rooms by means of modern data
logger, data writers or simple hygro- and thermometers.
However, the way and location of climate measurements are
often insufficient to reflect the real climate and to detect
different climatic zones in the building. In his book on micro-
climate in museums Camuffo (1998) illustrates the complexity
of climate monitoring that cannot just be monitored by
a single data logger in the middle of a museum room. The
influence of air stream through doors, warming by sunlight
and daily changes of temperature gradients as well as the
isolation and exposition of the building envelope have to be
considered as important factors. In fact, fungal growth mostly
happens between shelves with little aeration or near to
walls with temperatures below the dew point (Fig. 2E, F).
Micro-niches are often also created by wrapping of single
objects into plastic foils or extremely tight boxes not allowing
an exchange of air and vapour (Figs 1C, 2C, D).
The fungal micro-flora in museums is also influenced by
the atmospheric particular matter carrying carbonates,
minerals and others. Gysels et al. (2004) have shown in a study
on the Royal Museum of Fine Art in Antwerp that the indoor
aerosols were largely determined by the outdoor atmosphere
and the outdoor sources of organic and inorganic pollutants.
ants including polycyclic aromatic hydrocarbons (PAHs).
Therefore the fungal diversity on monuments in an urban
environmentwas foundtobe muchhigherthat in aruralenvi-
ronment of the climatic zone (Sterflinger and Prillinger, 2001).
Fungi as deteriorative agents of stone
From the biological point of view stone is an extreme environ-
ment poor of nutrients, with enormous changes of humidity,
with mechanical erosion due to wind and rain and high doses
of UV radiation. Nevertheless, stone is inhabited by fungi and
other microorganisms in all climate regions of the earth
(Sterflinger, 2005; Selbmann et al., 2005). Epilitic fungi – living
on the rock – and endolitic fungi – living inside of pores and
fissures – fungi play a major role in the weathering of monu-
ments made of rock. Fungi might be the most important endo-
lits on building stone, on mortar and on plaster because their
activity is high and they are extremely erosive (Sterflinger,
2000; Scheerer et al., 2009; Gadd, 2007). On monuments there
Role of fungi in deterioration of cultural heritage49
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Table 1 – Most frequent hyphomycetes in museums and on materials of objects of arta, b, c, d, e
Paintings: oil, water color, acrylicAlternaria sp.
Aspergillus flavus, Aspergillus sect. Niger, A. sydowii, A. versicolor
Cladosporium herbarum, C. cladosporioides
Eurotium chevalieri, E. rubrum
Penicillium chrysogenum, P. citrinum,
P. decumbens and many other species of the genus
Paper (laid-paper, wood pulp paper)
and cellulose textiles (cotton, linen)
Aspergillus clavatus, A. flavus, A. glaucus, A. terreus,
A. repens, A. ruber, A. fumigatus, A. ochraceus, A. nidulans, Aspergillus sect. Niger
Chaetomium globosum, C. elatum, C. indicum
Penicillium chrysogenum, P. funinculosum, P. pupurogenum,
P. rubrum, P. variabile, P. spinulosum, P. fellutatum, P. frequetans, P. citrinum
Trichoderma harzianum, T. viride
Parchment Cladosporium cladosporioides
(leather, wool, feathers, fur, hair)
Absidia glauca, A. cylindrospora, A. spinosa
Aspergillus sydowii, A. candidus, A. clavatus, A. carneus, A. foetidus,
A. flavus, A. fumigatus, and many other species of the genus
Cunninghamella echinulata, C. elegans
Penicillium brevicompactum, Penicillium chrysogenum and many other species of the genus
Archeological findings: bones, ceramicsArchaeological findings often carry a large load of spores, in case of
contamination the diversity on the objects reflects the diversity of the respective soil
The identification of the fungi was carried out based on morphology and/or sequencing of the ITS1, 5.8 S, ITS2 region with subsequent homology
search using the BLAST algorithm [http://www.ncbi.nlm.nih.gov/BLAST/].
a Unpublished data Sterflinger/ACBR.
b Mesquita et al. (2009).
c Meier and Petersen (2006).
d Blyskal (2009).
e Pangallo et al. (2009).
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are two major morphological and ecological groups of fungi
that are adapted to different environmental conditions. In
moderate or humid climates the fungal communities are
dominated by hyphomycetes including species of Alternaria,
Cladosporium, Epicoccum, Aureobasidium and Phoma. In arid
and semi-arid environments the fungal community shifts
towards black yeasts and microcolonial fungi. Black fungi
belonging to the genera Hortaea, Sarcinomyces, Coniosporium,
Capnobotryella, Exophiala and Trimmatostroma form small black
colonies on and inside the stone (Fig. 2A, B) and often occur in
close association with lichen (Sterflinger, 2005). Famous
monuments covered and deteriorated by fungi are the Acrop-
olis of Athens, marble monuments of the Crimea and the
antique temples of Delos (Diakumaku et al., 1995; Sterflinger,
2000). Due to their thick, melanized cell walls fungi resist
also chemical attack and cannot easily be killed by biocides
or other anti-microbial treatments. Black fungi dwell deep
inside granite, calcareous limestone and marble and deterio-
rate those stones. The phenomenon of biopitting – the forma-
tion of lesions in a size range of up to 2 cm in diameter and
depthonstone – iscausedby theblack fungi.Dueto thestrong
melanization of the cell walls stones inhabited by these fungi
appear spotty or are even completely covered by black layers.
In addition to outdoor environments black fungi are also
found on rock surfaces of caves and catacombs (Saarela
et al., 2004). The oldest and most precious objects suffering
from serious fungal invasions are rock art caves, especially
the Lascaux cave (Bastian and Alabouvette, 2009).
Fig. 2 – (A, B) Marble facade of the Celsus library in Ephesos (Turkey) with biopitting caused by microcolonial fungi. (C, D)
Fungal contamination of archeological findings and historical helmets due to storage in tight cardboard boxes. (E) Fungal
growth on textile tapestry on museum wall caused by wall temperature failing below the dew-point level. (F) The tight
assembly of racks for storage of paintings does not allow sufficient ventilation and thus increases the risk of fungal growth.
Role of fungi in deterioration of cultural heritage51
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4.Prevention and treatments
The three ‘‘ounces of prevention’’ against fungal contamina-
tion in museums are: climate control, frequent cleaning and
phenomenological monitoring. Climate concepts have to be
developed taking into account the individual architecture of
the museum. In spite of serious climate conditioning and
monitoring, unexpected water damage frequently happens
in old as well as new museum buildings. In this case the
hygienic situation – namely the amount of dust carrying
fungalspores– isdecisiveforthe dimension offungal contam-
ination. Objectsthat are heavily loaded with fungal spores will
easily be overgrown within a period of several days. However,
a surface carrying a limited amount of spores will be much
less affected. The air stream and air exchange in the rooms
as caused by the ventilation system, the number of visitors,
the action and air stream caused by the opening and closing
of doors has quite some impact on the load of fungal spores
carried inside a museum. New built storage rooms are now
provided with filter systems to avoid the invasion fungal
spores, plant pollen and dirt particles. Generally filter class
F5–F7 is used as final filter in air conditioning systems for
offices, sales rooms certain production plants and museums.
Frequent cleaning using vacuum cleaners equipped with
high efficiency particle absorbers (HEPA filters) is highly rec-
ommended to keep the spore load low.
Only recently important European museums have started
including the measurement of fungal spores in the air by air
sampling and the measurement of fungal spores on objects
and museum shelves by surface contacts (Fig. 3A, B). Based
on those measurements the hygienic status of the museum
can be determined, concepts for optimizing the hygienic
status and specific disaster management plans can be devel-
oped (Dicus, 2000; Barton and Wellheiser, 1985). Also the
implementation of quarantine rooms for contaminated
objects is increasing in museums and restoration studios.
In the process of restoration fungal spores and superficial
mycelia will be removed mechanically following a careful
investigation. The appropriate cleaning method is determined
by the chemical composition and strength of the material
itself and by the chemical quality of other than biogenic
patinas. Testing the viability of the fungi is crucial for making
the decision on further disinfection measures. Sampling of
fungal spores or small fragments of mycelium using a needle
(glass or surgical needle) is normally carried out directly in the
museums. Malt Extract Agar and Dichlorane Glycerol agar are
used routinely but a wide range of media might be used to test
the potential of the fungi to decompose the material: cellulose
agar, casein agar or stained indicator media (Pangallo et al.,
2009). On paper, pictures or photographs a defined part of
the surface can be sampled with a scalpel or a sterile swab
and being transferred to 0.9 % NaCl for transport to the lab
(Fig. 3C–E). The NaCl and the swab itself can be embedded in
suitable agar. In case the fungal flora turns out to be non-
viable, disinfection is not necessary.
For disinfection of a recent and progressive fungal damage
a limited range of physical and chemical methods are avail-
able (Allsopp et al., 2004), only the most common of which
can be discussed here: the most effective physical method
for killing fungi and their spores is the use of gamma radia-
tion. Most authors report that the doses has to be higher
than that normally used for bacteria and has to exceed
10–20 KGy for a complete killing of fungi (Nitte ´rus, 2000a).
However, gamma radiation might significantly affect the
chemical composition of the cellulose fibres and thus the
application has to be thoroughly considered. Chemical treat-
ments include liquid biocides and fumigation with gases
as methylbromide, ethylenebromide. The choice of the
appropriate biocide is somehow restricted by the European
Biocide directive (http://ec.europa.eu/environment/biocides/
index.htm). Albeit a variety of biocides is still available on
the market (Cooke, 2002) and in restoration three substance
groups are approved to be effective against fungi with none
or minimum irritation of the materials: (1) Products contain-
ing donors which slowly release formaldehyde. (2) Products
containing quaternary ammonium compounds with an
optimal chain length of C14–C16 (e.g. Metatin 5810-101, Neo
Desogen, Dimanin, Antimoos). These so-called ‘‘quats’’ are
rapid acting anti-microbials which are commonly regarded
as relatively environment friendly. Their efficacy is reduced
by high levels of salt or proteins. (3) Isothiazolinone, a more
recent biocide, was documented to be effective and even
preventive on paper objects. Treatments with different
phenols (e.g. thymol, cymol) gave good results in several
case studies but should not be regarded as general fungitoxic
solutions. Ethanol (70 %), the most common disinfectant used
in microbiology, can also have a good fungitoxic effect if the
time of application is at least 2–3 min. Mere spraying of
ethanol – as it is commonly carried out in restoration – is
insufficient (Nitte ´rus, 2000b).
5. Methods to study fungi on cultural heritage
Today, many sophisticated molecular techniques exist to
study the interaction of microbes and materials. However,
concerning the practice of conservation, consolidation and
prevention of biodeterioration the phenomenological analysis
of the objects using light and electron microscopy is the first
and most important step (De los Rios and Ascaso, 2005).
Mycological research on biodeterioration in museums or
monuments is still mainly based on classical cultivation
methods using a variety of standardized media as MEA,
DG18 and dichlorane rose bengale (DRBC) medium. In contrast
to bacteria for which it is generally accepted that cultivation
methods recover less than 1 % of the total present in environ-
mental samples, the recovery rate concerning fungi is
assumed to be more than 70 %. For this reason culture-based
approaches are still extremely useful in mycology.
Nevertheless, the use of modern molecular techniques for
the detection of fungi on and in materials of cultural heritage
will provide a deeper insight and understanding of fungal
community structures and their consequences for the mate-
rial. Recently, the Internal Transcribed Spacers (ITS regions),
which are nested in the nuclear rDNA repeat, have been
selected to investigate the fungal diversity of fungi on building
Rygiewicz, 2005). The ITS regions possess a high variation
52 K. Sterflinger
Author's personal copy
between taxonomically distinct fungal species and even
within the species. ITS based analysis of fungi on paper
from museums was carried out using Denaturing Gradient
Gel Electrophoresis (DGGE) (Michaelsen et al., 2006). For
studies of microbial communities colonizing artworks, DGGE
is the technique most often used (Portillo et al., 2008). An
example of a DGGE fingerprint from historical books is given
in Fig. 3F.
Fluorescence in-situ hybridization FISH has also been
applied in the field of conservation and restoration to study
bacteria, archaea and fungi involved in the biodeterioration
of surfaces (Urzı ` et al., 2003). Furthermore, the application of
FISH directly on adhesive tape strips added another advantage
to this non-destructive sampling method: the identification
‘‘in situ’’ of the microorganisms present on a given area,
withoutthe destructionof thevaluable surfaces and with little
Fig. 3 – (A) Air sampling in a restoration studio carried out with MD8 air sampler. (B) Sedimentation plates for monitoring in
a museum depot. (C) Sampling of medieval wall painting. (D) Sampling of photographs by swab within a marked area.
(E) Sampling of fungal mycelium on a historical book. (F) DGGE fingerprint of a fungal population of contaminated library
material showing remarkable differences between the books sampled (B1/B2) and between samples within books that were
taken from different macroscopically visible deterioration phenomena (lanes 37–40 and 28–34).
Role of fungi in deterioration of cultural heritage53
Author's personal copy
biofilm disturbance. Since FISH, using DNA probes, is often
hampered by rigid fungal cell walls, only recently peptide
nucleic acid (PNA) were applied for fluorescent in-situ detec-
tion of filamentous fungi (Teertstra et al., 2004). PNA probes
are synthetic DNA mimics, where the negatively charged
DNA backbone is replaced by a neutral polyamide backbone
(Stender et al., 2002). Due to this, PNA probes have better
binding features to complementary targets and penetrate
fungal cell walls more easily. This method could be a prom-
ising tool for specific detection and visualization of fungi on
and in materials.
Recent improvements in molecular studies have shown
the advantages of RNA-based molecular analyses. In an
RNA-based approach, not only the presence of a fungal
species but also its metabolic activity could be determined,
since the levels of RNA in a cell are proportional to the need
of that cell for synthesizing proteins required for metabolism.
RNA-based studies have been carried out to investigate the
yellow and grey colonizations on the walls of Altamira Cave,
Spain (Portillo et al., 2008). Also the analysis of the fungal pro-
teome (2-D electrophoresis or metaproteomics) can bring new
insights into the activity of fungi on art objects.
Fungi play a tremendous role in deterioration of our cultural
heritage. This holds true for museum objects as well as for
stone monuments in all climate zones of the earth. One
important reason why fungi are a great problem for conserva-
tion of cultural heritage is a lack of information and training
for restorers, curators and other museum personnel. There
is a high demand for mycologists and microbiologist able to
teach and to consult restorers and museum personnel. A basis
for this is a transdisciplinary approach in teaching and the
mycologists interest in material sciences and conservation.
This however is a big challenge for mycologists and microbiol-
ogist but also a very special field of working that offers inter-
esting insights into art and material sciences. It is worth
emphasizing the special role of fungal culture collections
where scientists have an extensive knowledge on fungal
taxonomy and ecology. Although mycologists knowledge
would be of tremendous value for museums, only very few
collections offer consulting in this field. To my knowledge
only one collection of those recognized by the World Federa-
tion of Culture Collections, namely the Austrian Center of Bio-
logical Resources and Applied Mycology (www.boku.ac.at/
acbr.html) offers a special service for museums and other
institutions working on care of monuments and cultural
I appreciate the Kunsthistorische Museum, the Theatre
Museum, the Essl collection, the Albertina Museum, the
museum of Ethnology (Vienna), the Academy of Fine Arts,
the University of Applied Arts, the Wien Museumand the Aus-
trian Ministry of Care of Monuments for their kind support
and for good collaboration.
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