Truffle cultivation and commercially harvested

native truffles

Dr. Shannon M. Berch

 PlacE! I Korea Forest Researc:h Institute
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Hosted by
• Korea Forest Research Institute
• Korean Forest Mushroom Society
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Supported by
• Korea Forest Service
• Pochun Mushroom Development Co. Ltd.
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 Truffle cultivation and commercially harvested
native truffles

Dr. Shannon M. Berch

British Columbia Ministry of Environment, Environmental Sustainability and Strategic

Policy Division, Ecosystems Protection & Sustainability Branch, Victoria, BC,
Canada; [email protected]

Abstract
Truffles (ftue and false) are fruiting bodies of ectomycorrhizal fungi and some of them
produce appealing aromas, are recreationally and commercially harvested, and even
.. cultivated. Until recently, commercial truffles have all been Mediterranean in distribution
but .some of these species are now cultivated around the world and other native species
are being collected and marketed. While cultivation of black truffles can be complicated
by horticultural challenges, production of other species appears to be less problematic. The
potential for the discovery and commercialization of novel native truffles is good, but
only if trained dogs are used for exploration and harvesting can this potential be
sustainably and ethically realized.

Introduction

Around the world, ftuffles are harvested primarily because of their enticing aromas.
They are generally considered as a delicacy and a luxury. All ftuffles (true and false) are
the fruiting bodies of ectomycorrhizal fungi and therefore are associated with living host
plants, usually ft'ees or shrubs. In Europe, most commercial truffles are members of the
genus Tuber but in dry ecosystems of Africa other genera dominate, e.g. Telfezia and
Tirmania.

85
 Because of their ectomyconhizal nature, some truffles can be cultivated in orchards
with their plant hosts. Both wild-harvested and cultivated truffles are highly sought-after
•
and some species command extraordinary prices; Hall et al. (2007) quote retail prices in
2005 for Italian white uuffle (Tuber magnatum) of up to €6,000 per kilogram and for
Perigord black truffle (Tuber melanosporum) of €3,000. Market prices taken from Bonito
et al. (2013) indicate that prices per kilogram in 2009 in the USA ranged from $110
USD for Tuber indicum to $5060 for Tuber magnatum' (Table 1). Charity auctions in
Alba, northern Italy, often catch media interest as prices paid for Italian white truffles can
be astronomical. In 2010, for instance, a giant (900 gm) white truffle was sold to Jeannie
Cho Lee, a South Korean wine critic living in Hong Kong, for $144,000.
These high prices make the prospect of cultivating truffles velY attractive and have led
to a great deal of research and development in the truffle industry globally. Unfortunately,
the hope of big financial returns has also fostered an air of secrecy and even deception
in parts of the truffle indusuy. Despite some highly rewarding success stories in truffle
cultivation, many truffle orchards fail to produce commercial amounts of truffles for a
variety of reasons, some of which will be mentioned below.

Table 1. Economically important truffle species determined by market pnces (USD).

Adapted from Table S2 in Bonito et al. 2013 and ii-om Marte et al. 2012.

Tuber ilJdicum $110

Tuber canaliculatum $220 North America
Tuber gibbosum $220 North America
Tuber lyonii $220 North America
Tuber oregonense $220 North America

Tuber borchii $440 Europe

Tuber brumale $660 Europe
Tuber macrosporum $660 Europe
Tuber melanosporum $1760 Europe
Tuber magnatum $5060 Europe

Desert nuffles* $26 - $330 Europe, Africa

*from Marte et al. 2012.
 are ascomycetes, primarily in the genus Tubel~ and false truffles are
the majority of commercial huffles are members of the genus Tuber.
,pnito et al (20l0a) predicted that global Tuber species dchness would be a minimum of
',80-230 species based on their meta-analysis of Tuber ITS rDNA sequences. Because of
mall size, disagreeable odor, or rarity, not all species of Tuber are considered desirable
pr personal or commercial harvest. Among the approximately 350 hypogeous ftmgi in the
. acific Northwest region of USA including hue truffles and false ·truffles (Trappe et al.
only a small handful of species are considered edible and choice. In their
omprehensive treatlnent of truffles and truffle cultivation, Hall et al. (2007) outline the
or so huffle and false tmffle species of commercial importance globally.
i{\The commercially impOltant MeditelTanean true truffles include some species that are
!tpw cultivated well outside of their native ranges: Perigord black huffle (Tuber
m~lanosporum), summer huffle (Tuber aestivum), and, bianchetto (Tuber borchii). In China,
number of tmffle species are harvested; among them Tuber indicum is the most
wpoltant. Growing interest in native tmffles in North America has provided a market for
regon white truffles (primarily Tuber gibbosum and Tuber oregonense) and the pecan
ff1e (Tuber lyonii).
dry ecosystems of the Carpathian Basin, Middle East, MeditelTanean Basin, and
orth Africa, other hypogeous ascomycetes are harvested including Mattirolomyces
etfezioides, Tel!ezia arenaria, Kalaharituber pfeilii, Tirmania nivea, Delastria rosea, and
{eoa lefebvri (Hall et al. 2007). Desert tmffles tend not to have the pungent aromas of
ffber species so rather than being harvested with the help of trained dogs, the
haracteristic cracks that form in the soil directly above them are recognized by
,l'pedenced harvesters. Considered a treat by some, desert huffles have also traditionally
Jlpplied much needed nutrition to people living in harsh, add environments.
..Among the false huffles, Rhizopogon luteolus, R. piceus and R. roseolus (~ R.
ibescens) are considered edible (Boa 2004), and in New Zealand R. roseolus is being
cliltivated (Visnovsky et al. 2010).

87
 Commercially harvested wild truffles

Europe is home to a number of commercially import truffle speCIes (Hall et al. 2007).
In the black tlUffle group are Tuber melanosporum, Tuber aestivum, Tuber brumale, T.
mesentericum, and T. macrosporum. The white ttuffles include Tuber magnatum, T. borchii,'
and T dlyophilum. To the real tlUffle connoissem who knows where and when to look,
many more species are available and Alessandra Zambonelli (personal communication,'
2011) can enjoy ttuffles evelY weekend of the year that are wild-harvested within
reasonable driving distance of her home.
Commercial Asian ttuffles of the black ttuffle group are still somewhat in taxonomic
flux (Zhang et al. 2005, Wang et al. 2006, Bonito et al. 20lOa) but Tuber indicum is
thought to be the most important species (Hall et al. 2007). Shin et al. (1995) reported
,
finding Tuber aestivum in Gyerong-san National Park, South Korea, under Quercus, bul
re-examination of this collection using molecular techniques is warranted given the currenf
state of taxonomic flux. There has been a recent proliferation of new Tuber species
descriptions from China (Fan and Cao 2012, Fan et al. 20 11a, Fan et al. 2011b, Fan et
ai. 2012, Zhang et ai, 2012), so it seems reasonable to anticipate that the hypogeon'
fungi of Asia are much more diverse that it may cmrently appear (Wang 2012). How
many Qf them will appeal to the palate remains to be seen.
White tlUffles, black tlUffles (Leucangium carthusianum), and brown tlUffles (Kalapuy'
brunnea) are commercially harvested from the wild on the west coast of NOlth America·
Although all of these commercial ttuffles fluit in young Douglas-fir (Pseudotsui'
menziesii) stands, the Oregon white truffles in pmticular fruit abundantly in over-groW
Douglas-fir Christmas tt'ee farms that were planted in agricultural fields adjacent to nativ'
Douglas-fir forests (Pilz et al 2009). Although nmselY inoculations of Douglas-fir wi
Tuber oregonense, T' gibbosum, and L. carthusianum have resulted in well-colonize
seedlings (Lefevre 2012), attempts to establish orchards have been limited by the cost!'
nurSeIy production. According to Lefevre (2012), three eastem North American
have commercial potential: Tuber lyonii, Tuber canaliculatum, and Imaia gigantea. Tub
lyonii, the pecan ttuffle, is harvested from native forests but also fi'om orchards of Cal
illinoinensis where it seems to thrive under the conditions of irrigation, fertilization a~,
chemical weed control practiced in pecan production. Tuber canaliculatum and Imaf
gigantea are known to have good culinmy attributes but are relatively seldom encounterc
in the'wild and not yet cultivated.
 esert truffles have a long history of use in many arid and semi-arid parts of the
editelTanean basin and Africa (Morte et al. 2012). In contrast to the other tlUffles which
iroarily associate with trees and some ShlUbs, desert ttuffles form myconhizas with
robers of the Cistaceae including both perennial and annual Helianthemum species.

,Tuber me/anosporum is now cultivated in all of the continents with arable land
eluding throughout Europe (Hall et al. 2007); Africa (Morocco, Abdelaziz Laqbaqbi,
l'sonal communication, 2011); Asia (China, Wang and Hall 2004); Australia (Australia
d, New Zealand, Hall et al. 2007); North America (USA, Lefevre 2012); and South
'erica (Chile, Cordero et al. 2011). Tuber aestivum and Tuber borchii are also now
qely cultivated and 'superb' bianchetto truffles are being harvested in a trial plantation
Zealand (Alexis Guerin-Laguette, Plant and Food Research, personal
July 2013). These two species have broad host ranges, wide ecological
established markets (Benucci et al. 2012). However, despite repeated
fteropts, the most highly priced tlUffle, Tuber magnatum or the Italian white tlUffle, has
.Qt yet been produced in cultivation. Cultivation of these Mediterranean Tuber species
'ually involves establislnnent of orchards of tmffle-inoculated oak (Quercus) or hazelnut
diy/us) trees.
Since 1999, TerJezia clavefyi has been cultivated in Spain, Israel, Abu Dhabi and
gentina on a variety of Helianthemum species (MOlie et al. 2012) and recently the
.1tivation of Tefjezia bouderi on Helianthemum sessiliflorum in Tunisia has been reported
,
lama et al. 2010).
Hf\"
fRhizopogon roseo/us (synonym Rhizopogon rubescens Tul.) or shoro has been cultivated
!l
, Pinus )'adiata in New Zealand for the Japanese shoro market (Wang et al. 2012).
:j~novsky et al. (2010) report~d that the shoro crop from New Zealand was "deemed
H~~itable due to consumer sensitivity in Japan to the origin of the products and doubts
&rounding the authenticity of the fiuiting bodies as Japanese shoro". They detennined
ii!
'at shoro produced in New Zealand was more closely related to collections from North
.) erica than it is to shoro from Japan, a finding that is consistent with the originally
cidental and later intentional introduction of Rhizopogon raseo/us to New Zealand for the
x!'),

'Itivation of Pinus radiata. Now however, Japanese isolates of this fungus are being

89
 cultivated in New Zealand and they may prove to be more acceptable to the Japanese
market (Alexis Guerin-Laguette, Plant and Food Research, personal communication, July
2013).

Recent achievements in truffle cultivation in North America

Perigord black tlUffles have been harvested from truffle orchards in California, North
Carolina, and Tennessee (USA) for a number of years although the amount of commerciai
product is still quite limited. A tlUffiere in the Willamette Valley recently provided
Oregon's first Perigord black truffles
(http://www.truffletree.comlfirst-cultivated-french-black-truffle-found-in-oregons-willamette-v
alley!).
Three Perigord black truffles were harvested by trained dog from a truffle orchard in
the Lower Fraser Valley in Abbotsford, British Columbia, Canada in March
(http://www.bctlUffles.caIHome/NewsDetaiI/26). These ltuffles represent the
independentIy-confllmed production in a tlUffle orchard in Canada.
In FebrualY of 2012, North America's first bianchetto huffles were harvested in Idaho,
USA (http://www.truffletree.comlnew-idaho-tuber!).

Challenges

The major challenge is that despite a global effort to establish tlUffle orchards over the
.
past few decades, truffle production continues to decline and Hall et aI. (2007) provided
an anay of reasons why many truffle orchards fail to produce. The original motivation
for establishing a tlUffle orchard for many is not actually to produce huffles, though thai
. "1/
would be a pleasant bonus, but to take advantage of tax breaks and grants or to skiM:
around regulations intended to product agricultural land £i'om development. Many tlUffi~r~J.
are established under sub-optimal conditions of climate, soil, or plant/fungus quality ~nd
many are not properly managed after establishment.
Cultivating exotic species in novel environments creates the risk that exotic species rna
invade native ecosystems and cause problems. There are many examples of this happenin
" (1
when plants (e.g. trees and slmlbs, Richardson and Rejmanek 2011) and animals (Jeschke
and Strayer 2005) are purposefully introduced; problems created by non-native fungi are
often the result of accidental rather than intentional introductions, e.g. Ophiostoma
(Dutch elm disease, Temple et al. 2006) and Amanita phalloides (death cap
Pringle et al. 2009) in North America. Truffles such as Rhizopogon species
vere inadvertently introduced into New Zealand on the roots of Pinus radiata seedlings
(Visnovsky et al. 20 I0) decades before they were intentionally cultivated there.
'With the intentional introduction of high value truffle species into novel ecosystems
comes the possibility, perhaps even likelihood, of unintended introdllction of other truffle
§pecies. Within the black tr'uffle group, Tuber melanosporum has the greatest commercial
alue which makes it the species most often introduced, however, other lower value but
orphologically similar black truffle species such as Tuber brumale and Tuber indicum
ave been reported from Perigord black truffle orchards in Australia, Italy, New Zealand,
nd United States of America (USA) (respectively, Linde and Selmes 2012, Murat et al.
c008, Ho et al. 2008, Bonito et al 20 IOb), presumably as a result of mistaken inclusion
l,)f these species in the sporocarp-based inoculant used in commercial nurseries to produce
truffle-colonized seedlings for out-planting in truffle orchards. In fact, Bonito et al (2010a)
pllve provided evidence suggesting the presumably accidental introduction of many other
tmffles species (T foetidum, T menseri, T rapaeodorum, T levissimum, T maculatum, T
II/um, T seperans, T californicum, as well as a number of un-named Tuber species)
mong Europe, North America, New Zealand, and Argentina.
While import regulations may be intended to assure the quality and identity of truffle
roducts, in reality it would be impossible to stop the movement of undesirable or
prohibited truffle species. Collections of Perigord black ttuffles being imported into the
SA, for instance, have been found to contain other black truffle species (Jim Trappe
nd Charles Lefevre, personal communications 2012). Border agents cannot be expected to
1'. able to identifY tlUffle species when truffle exporters, who should be able to do so,
., arly make mistakes. Murat et al. (2008) reported that a specific regulation established
1985 and modified in 1991 prohibits the importation of Tuber indicum into Italy; sadly,
.llt regulation failed to keep it out of a Tuber melanosporum orchard. Similarly, New
ealand's biosecurity standard for importation of fi'esh 01' frozen ttuffles (New Zealand
inistry of Agriculture and Forestry 1993, amended 2011) permits the impOliation of four
only (Tuber aestivum, T borchii, T magnatum, and T melanosporum). When Tuber
was· detected in a quarantine facility in New Zealand in 2006, an eradication

91
 program was launched; when T. brumale was subsequently found on the roots of trees
planted much earlier, the program was terminated (Ho et aI. 2008).
Seedlings in commercial nurseries are inoculated with slurries (Hall and ZamboI)elIi
2012). These SIU11'ies are often made from ttuffles with imperfections that make them
more affordable than marketplace specimens and from pieces of truffles. It is both
difficult and time consuming to distinguish among similar truffle species by examining
whole truffles or pieces of tlUffle that are impOlted as inoculum. TlUffle experts now rely
on DNA sequencing to confirm the identity of huffles. Truffle expOlters with the velY
best of intentions cannot DNA sequence evelY piece of truffle that is exported; truffle
exporters who are seduced by the possibility of making even more profit by substituting
low value for high value truffle species might slip a few Tuber brumale or Tuber indicum
into a shipment purported to be pure Tuber melanosporul1l. Reputable connnercial growers
of inoculated seedlings are obliged to DNA check each and every huffle or piece of
truffle used as inoculant and to record to which inoculant lot each seedling belongs so
that any contamination can be traced from source to shipment to orchard. If even a tiny
shard of low value truffle makes it through the checking process, the mature spores in it
could contaminate an entire lot of host tree seedlings.
Other quality control criteria include the extent of colonization by the desired truffle
speCles, the absence or minimization of colonization by undesired ECM fungi, size and
vigour of the seedling (Bonet et aI. 2009, Fischer and Colinas 1996), and in North
America the resistance of the C01ylus avellana variety to a fungal blight Anisogramma
anomala. In Europe, growers are discouraged from planting COIylus avellana not because
of this blight fungus but because hazelnut is more likely to host Tuber brumale than are
oaks.
On the surface, it seems that another approach to quality control would be to set
criteria for c~ercial nurseries. Italy, France and Spain all have protocols for
certification of truffle-inoculated nurselY trees and the Austt'alian Truffle Growers
Association is considering a similar approach (Carter 2011). All of these approaches raise
the bar by demanding conscious commitment to high quality but not all ECM tips or all
seedlings from a nursery can be examined before shipment so the possibility. of
contamination, even if very low, will always exist.
Duffles are traditionally harvested using a trained dog or a pig because these animals
have keen noses and will mark only mature truffles with good commercial value. In
China, much of the truffle harvest is raked and this may have lead to a decline in
 volume harvested over the years. According to Murat et ai. (2008), before 1993 over 20
tons of Tuber indicum were halvested in Huidong County alone while in 2003 less than 5
tons were halvested from the same area. In the Pacific NOIthwest, USA, ttuffles were for
awhile hatvested almost exclusively by people with rakes which led to the collection of
many immature truffles with poor aroma that sold for rather low prices (Lefevre 2010);
education and proper training have contributed to the increase in the number of Oregon
white, black and brown tt'uffles that are now dog-harvested.

Opportunities

While cultivation of Tuber melanosporum seems fraught with difficulties related to

contamination of tmffieres by morphologically-similar but lower-value species, cultivation
of Tuber aestivum and Tuber borchii appears to be more promising. Although the potential
returns are lower when compared to successful cultivation of Tuber melanosporum, many
growers may prefer the lower risk associated with summer truffle and bianchetto.
The imposition of proper hatvesting techniques using trained dogs in North America
(Lefevre 2012) and China (Wang 2012) wiII without a doubt improve the quality and
thereby'the value of their native truffles.
Because they wiII help ensure that mature truffles are found and available to tmffle
taxonomists, tt'ained tmffle dogs will also be very useful in expanding what we know of
the truffle mycota of the parts of the world where little is currently known. The
systematic application of DNA techniques to ttuffle taxonomy III combination with more
tt'aditional morphological approaches will guarantee that the new species being found can
be integrated into our global knowledge of these fungi.
Additionally, phylogenetic analysis could be used to assess the potential edibility and
commercial potential of new species being found. Wang (2012) points out that two
Chinese truffle species could have commercial potential because they are closely related to
Tuber borchii.
The native Oregon truffle industty developed in part because the ttuffles of Oregon
have been extensively collected and studied by Dr. Jim Trappe, his students, and by
members of the North American Truffling Society (NATS). By organizing regular truffle
forays, producing guidelines to protect the truffle enviromnent and minimize harvest of
immature truffles, promoting truffle dog tt'aining, and closely collaborating with truffle

93
taxonomists, NATS has fulfilled its mission "to enhance the scientific knowledge of NOlih
American truffles and tl1lffle-like fungi". Application of this model elsewhere would do a
lot to improve our knowledge of ttuffles. Whether we have enough truffle tajmnomists or
jobs for students of truffle taxonomy to support local ttuffling groups is a question that
remains to be addressed.

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