ArticlePDF Available

Abstract and Figures

Blueberries belong to the genus Vaccinium, a widespread genus with more than 200 species of woody plants. In Northern Europe, the European blueberry (EB), also called bilberry, is one of the most important wild berries. EB (Vaccinium myrtillus) is very demanded by the processing industry, due to its delicious taste and high dietary value. However, to our knowledge there has been made no efforts of domestication of the species, and it is still harvested in forest fields without any cultivation. The successful management of the sweet lowbush blueberry (V. angustifolium), which in many ways is similar to the EB, suggests that there are opportunities to increase yield and decrease the significant yearly variation in EB yield, by practices including fertilization, irrigation, cutting trees, and weed control. The fruit yield in wild stands of EB is very variable, but the potential is probably close to 2 tons per hectare. Results from literature on growth of the EB, development and ecology are discussed in relation to possibilities for domestication.
Content may be subject to copyright.
Received: 21 April, 2010. Accepted: 29 July, 2010. Invited Review
The European Journal of Plant Science and Biotechnology ©2011 Global Science Books
The European Blueberry (Vaccinium myrtillus L.)
and the Potential for Cultivation. A Review
Rolf Nestby1* David Percival2 Inger Martinussen3 Nina Opstad4 Jens Rohloff5
1 Norwegian Institute for Agricultural and Environmental Research, Grassland and Landscape Division, Kvithamar, 7500 Stjørdal, Norway
2 Department of Environmental Sciences, Nova Scotia Agricultural College, Truro, Nova Scotia, Canada
3 Norwegian Institute for Agricultural and Environmental Research, Arctic Agriculture and Land Use Division, Box 2284, 9269 Tromsø, Norway
4 Norwegian Institute for Agricultural and Environmental Research, Arable Crops Division, Kapp, Norway
5 The Plant Biocentre (PBC), Department of Biology, Norwegian University of Science and Technology (NTNU), Trondheim, Norway
Corresponding author: *
Blueberries belong to the genus Vaccinium, a widespread genus with more than 200 species of woody plants. In Northern Europe, the
European blueberry (EB), also called bilberry, is one of the most important wild berries. EB (Vaccinium myrtillus) is very demanded by
the processing industry, due to its delicious taste and high dietary value. However, to our knowledge there has been made no efforts of
domestication of the species, and it is still harvested in forest fields without any cultivation. The successful management of the sweet
lowbush blueberry (V. angustifolium), which in many ways is similar to the EB, suggests that there are opportunities to increase yield and
decrease the significant yearly variation in EB yield, by practices including fertilization, irrigation, cutting trees, and weed control. The
fruit yield in wild stands of EB is very variable, but the potential is probably close to 2 tons per hectare. Results from literature on growth
of the EB, development and ecology are discussed in relation to possibilities for domestication.
Keywords: aerial shoot, buds, cultivation, flowers, fruits, rhizome
INTRODUCTION.......................................................................................................................................................................................... 5
VACCINIUM – SYSTEMATICS.................................................................................................................................................................... 7
PLANT CHARACTERISTICS...................................................................................................................................................................... 7
Rhizomes and roots ................................................................................................................................................................................... 7
Aerial shoots.............................................................................................................................................................................................. 8
BUD DEVELOPMENT AND FLOWER INDUCTION ............................................................................................................................... 8
POLLINATION AND FRUIT DEVELOPMENT.......................................................................................................................................... 8
FRUIT CHARACTERISTICS....................................................................................................................................................................... 9
POTENTIAL OF ENHANCING PLANT DEVELOPMENT AND FRUIT YIELD BY MANAGEMENT ................................................. 9
Half cultivation.......................................................................................................................................................................................... 9
Forest cutting........................................................................................................................................................................................... 10
Climate .................................................................................................................................................................................................... 10
Soil and nutrition..................................................................................................................................................................................... 11
Insects...................................................................................................................................................................................................... 11
Diseases................................................................................................................................................................................................... 11
Weeds ...................................................................................................................................................................................................... 12
Browsing animals and birds feeding on the berries ................................................................................................................................. 12
Cultivation on farm land.......................................................................................................................................................................... 12
REPRODUCTION....................................................................................................................................................................................... 12
Seeds........................................................................................................................................................................................................ 12
Vegetative propagation............................................................................................................................................................................ 13
HARVEST AND DELIVERY...................................................................................................................................................................... 13
CONCLUSION............................................................................................................................................................................................ 13
REFERENCES............................................................................................................................................................................................. 14
The European blueberry (EB) is a deciduous woody dwarf
shrub, typical and abundant especially in spruce- and pine-
dominated heath forests of medium fertility in the northern
hemisphere (Fig. 1).
EB also grows in marginal types of forests, and above
the tree limit up to high altitudes. Typically, it grows on
areas of better-drained acid soils. The tiller (individual
shoot from the rhizome) develops into an axis (a system of
annual shoot increments formed by the branching of the
original tiller or subsequently, by the branching of a shoot
arising from the lower part of this; of limited longevity)
from approximately 5 to 90 cm high depending on climate
and nutrient availability (Flower-Ellis 1971). The bush,
consisting of abundant tillers, is perennial and deciduous,
and leaves are notably bright green, 1-3 cm long, slightly
toothed and not leathery. Tillers and leaves are hairless and
The European Journal of Plant Science and Biotechnology 5 (Special Issue 1), 5-16 ©2011 Global Science Books
can by this be distinguished from other members of the
heather family or any other moorland or mountain shrub
even without flowers and fruit. It plays a significant role for
a number of other species, particularly of birds and mam-
mals, and its abundance has been used as an indicator of
biodiversity within the forests. The EB has long had an im-
portant role in human cultures; the nutrient rich berries
providing food, and used as a herbal remedy for digestive
problems, diabetes and to strengthen capillaries in the cir-
culatory system. The juice of the berries has also been used
as a dye, providing a blue colour for both linen and paper
(Featherstone 2002).
Cultivation of Vaccinium spp. is not a new approach
within horticulture. Blueberry cultivation is thought to have
started when Native Americans burned wild stands of low-
bush blueberries (V. angustifolium Ait. and V. myrtilloides
Michx.) to tend them and to increase production (Strik
2006). More advanced blueberry domestication began in
New Hampshire in 1908 by the United States Department
of Agriculture (Janes and Percival 2003). Since the 20th cen-
tury American blueberry species and varieties have been
spread to Europe (Nyström 1932; Kühn and Vang-Petersen
1991; Gough 1996; Hjalmarsson 2006). Some of the know-
ledge achieved from the very successful domestication of
highbush blueberry species, could be transferred to the pot-
ential cultivation of the European blueberry (EB). However,
the approaches and associated technologies developed with
the sweet, lowbush blueberry (V. angustifolium) in North
America, may present an increased likelihood of success.
This plant is comparable in many ways to V. myrtillus, with
similar growth, morphological characteristics (e.g., exten-
sive root and rhizome system), berry size, coloration and
good winter hardiness.
Domestication of the lowbush blueberry began in Maine
in the 19th century, and the blueberry industry has since
undergone a significant increase. In 1990, the industry had
over 1000 producers and the lowbush blueberry acreage and
production were respectively 113 000 ha (Kinsman 1993)
and 12 700 tons, compared to 477 tons in 1951. In 2008 the
production was increased further and was approximately
113 000 tonnes (Percival 2010 pers. comm.). The size of
individual producer operations varies from one to over 10
000 ha. Most commercial fields have been developed from
abandoned farmland and woodlands that were deemed to
have substantial lowbush blueberry plants present after tree
removal. Although lowbush blueberry cultivars have been
developed, the crop in Maine, the Canadian Maritimes, and
Quebec is almost entirely produced by genetically diverse
wild clones (Aalders et al. 1979; Hall 1979; Chen et al.
1998). To make a similar success history in an industry
based on the EB may be difficult, as EB are a dwarf shrub
that produce single or paired berries on the bush instead of
clusters, and consequently have relatively low yields. How-
ever, the demand for quality fruit of EB is strong due to a
high content of valuable biological compounds (Halvorsen
et al. 2002; Riihinen et al. 2008).
The EB is a calcifuge plant that is circumboreal in dis-
tribution. It occurs in Europe, the northern parts of Asia,
including Japan, and also in Greenland (Fig. 1) where it is
thought to be introduced from Europe, possibly in Viking
times (Featherstone 2002). In Europe it is still harvested in
rural areas by the local population (Pardo de Santayana et al.
2005; Konovalchuk and Konovalchuk 2006; Coudun and
Gegout 2007) for household consumption and recreational
purposes (Kangas and Markkanen 2001), and is one of the
economically most important wild berry species in Finland,
Norway and Sweden (Kangas 2001). For the industry in
Sweden, Finland and some Central and East European
countries EB are hand harvested by combing the plants in
cut forestland. Intensive forestry with clear-cuttings and soil
preparation has been observed to give a more even distribu-
tion of the different age classes and to decrease the cover-
age of EB (Kardell and Eriksson 1995; Miina et al. 2009).
On the contrary, Nielsen et al. (2007) found that EB pro-
duction and growth decreased with increased forest maturity.
Therefore, for a stable and possibly increasing supply of
berries to the EB industry, there is a need to develop an
improved and more scientifically based production system
Fig. 1 Distribution of EB in the Northern hemisphere (Hultén and Fries 1986), with the kind permission of Sven Koeltz (Dnr 70-79/1998).
European blueberry domestication. Nestby et al.
in which consistent, high yielding crops of EB with excel-
lent quality attributes are produced with manageable input
costs (e.g. harvesting costs, cost for transportation and
logistics). Several aspects need to be taken into considera-
tion in developing an improved production system, either
growing EB on abandoned farmland, or by half-cultivation
on cut forestland.
To our knowledge, there have been no prior systematic
examinations of how to domesticate the EB by half-cul-
tivating forest fields or by growing on farm land. There are
however, numerous publications discussing the biology and
ecology of EB in forest ecosystems, and how its distribution
is affected by natural disturbances and forest cultivation.
The objective of this paper is to present an overview of rele-
vant work dealing with EB, and relate it to possibilities for
cultivation. The sweet lowbush blueberry will be the bench-
mark model used since it is highly domesticated and has
been thoroughly examined for reactions to environmental
conditions (Kinsman 1993; Jeliazkova and Percival 2003b).
The genus Vaccinium consists of woody, perennial shrubs in
the heath family (Ericaceae) and contains, among other spe-
cies, both the European and the American blueberry (Tab l e
1). Although the EB is often referred to as the bilberry, this
includes several species of low-growing shrubs in the genus
Vaccinium that bears fruit. The term European blueberry
(EB) will be used throughout this article since it points
directly to V. myrtillus and defines it as a blue berry which
is mirrored in most European languages. Other names are
blåbær (Scandinavian), blaeberry (Britain), whortleberry
(Britain), whinberry (or winberry, Britain), wimberry, myr-
tle blueberry, heidelbeere (Germany), blaubeere (Germany),
fraughan (Ireland), myrtilli (Italian) and other names regi-
onally (Lid 1963; Tirmenstein 1990; Featherstone 2002;
Wikipedia 2009a). The European and the American blue-
berry species belong to different sections, and while the EB
belongs to the section Myrtillus together with 11 other spe-
cies of minor importance, the American blueberries belong
to the section Cyanococcus (Ta b l e 1). The EB is a shrub
varying in size from approximately 5 to 90 cm, and chro-
mosome number is 2n=24 (Hagerup 1928; Hedberg and
Hedberg 1961; Tirmenstein 1990; Vander Kloet and Dickin-
son 1999; Featherstone 2002). The American blueberries
vary in size from approximately 10 to 400 cm with the
smaller species known as "lowbush blueberries" (typically
less than 1 m in height) and the larger species as "highbush
blueberries". The flowers of both sections are bell-shaped,
white, pale pink or red, sometimes tinged greenish. How-
ever, while the EB grow one (rarely two) flower from a leaf
axil bud, the American blueberries can produce an inflores-
cence containing a cluster of berries from a leaf axil bud.
Highbush and lowbush blueberries develop relatively large
clusters of berries mainly on one-year-old wood but also
some on two and three year old wood, producing greater
yields than huckleberries. Western huckleberries are in dif-
ferent taxonomic sections (Myrtillus, Vaccinium, and Pyxo-
thamnus) than highbush and lowbush blueberries (Cyano-
coccus). Sections Vaccinium and Pyxothamnus each contain
one species. Some species are found not only in North
America, but also in Europe, Asia, and Greenland (Barney
2008). The two species V. uliginosum (Bog bilberry) and V.
vitis-idaeus (lingonberry) grow frequently under the same
environmental conditions in the north of Europe as EB, and
belong to the sections Vaccinium and Vitis-idaea respec-
Rhizomes and roots
The characteristics and growth pattern of EB are well docu-
mented in the work of Flower-Ellis (1971). EB growth pat-
tern is rhizomatous and it usual forms open colonies or pat-
ches in the vegetation that can be as large as 15 m in dia-
meter (Ritchie 1956). A characteristic form of the bilberry
stand is that of an irregular, approximately circular patch,
with an increase in rhizome age from the distal of the stand
to the centre (Flower-Ellis 1971). The growth direction of
rhizomes is mainly centrifugal, but soil factors strongly
affect rhizome growth directions so density and rhizome
growth and diameter growth is not necessarily correlated.
Rhizomes up to 5.5 m long have been found, with an ave-
rage length of 2.0 m (Albert et al. 2003), and one single
clone may occupy several square meters (Flower-Ellis
1971). The branching of the rhizome is sympodial and the
aerial shoot disposition is reflected by this. Typically, from
their originating point two rhizomes extend 20-30 cm
underground without branching before dividing, ideally into
two, occasionally into three or more, of which the middle
one is the most vigorous. This middle rhizome turns up-
wards and develops a green shoot or continues its growth
underground. After a further 20-30 cm it divides into a
group of shoots which corresponds morphologically with
the green aerial shoot. Thus the rhizome in EB ends in a
green shoot or in a corresponding group of rhizomes. Fur-
ther growth is continued by two side branches. Along a seg-
ment of rhizome, aerial shoots tend to arise at intervals of
20-30 cm (Flower-Ellis 1971). EB rhizomes can live for up
to 34 years, although rhizomes older than 15 years rarely
produce new aerial shots or rhizomes. A concern that has
sometimes surfaced in wild blueberry growth is stagnation
of growth caused by a lack of rejuvenation of the rhizome
system (Percival 2010 pers. comm.).
The quantity and size of clones (one rhizome and
shoots/axis originating from it) may vary significantly, even
within the same population (Albert et al. 2004). Nutrient
mobilization and reallocation typically occurs in the autumn
with nutrients mowing from the leaves to the rhizomes, and
this is the main time for rhizome growth, along with the
spring (Featherstone 2002). New roots are mainly found in
late summer and in autumn, and outside this period no im-
portant root formation occurs, even in late spring and early
summer, when new aerial vegetation develop and flowering
take place (Bonfante-Fasolo et al. 1981). However, given
Table 1 The systematic link between European and American blueberries.
Order Ericales
Family Ericaceae
Subfamily Vaccinioideae
Tribe Vaccinieae
Genus Vaccinium Section Important species of section
Batodendron, Brachyceratium, Bracteata, Calcicolus, Ciliata,
Cinctosandra, Conchophyllum, Cyanococcus, Eococcus, Epigynium,
Hemimyrtillus, Herpothamnus, Myrtillus, Neurodesia, Oarianthe,
Oxycoccoides, Oxycoccus, Pachyanthum, Polycodium,
Pyxothamnus, Vaccinium, Vitis-idaea.
V. myrtillus
V. corymbosum
V. angustifolium
V. myrtilloides
V. virgatum (ashii)
V. darrowi
The European Journal of Plant Science and Biotechnology 5 (Special Issue 1), 5-16 ©2011 Global Science Books
the symbiotic nature of the fine roots with mycorrhizal
fungi, there is probably always some level of new root
growth and development occurring (Percival 2010 pers.
Aerial shoots
Aerial shots of EB arising from the same rhizome system
have similar characteristics and are referred to as ramets
that together with the rhizome, originate from and form a
blueberry clone (Fig. 2). The resulting entity of ramets/axes
of one clone, form a bush (Flower-Ellis 1971).
However, EB has high morphological plasticity, and
growth habit in terms of e.g. branching can differ greatly
between habitats as a response to environmental differences
(Tolvanen 1994). EB aerial shoots grow more vertically and
have lower branching angles in open habitats compared
with the forest. The stems are angled and woody, and the
continuous rejuvenation from the abundant bud bank into
many annual shoot generations, leads to a complex shoot
age structure within ramets. The growth form of EB clones
is phalanx rather than guerrilla (Albert et al. 2004), imply-
ing that clones produce short internodes and closely packed
ramets, and to a lesser degree long internodes and widely
spaced ramets. Average ramet age is also affected by its
habitat, and increases with forest maturity in older stands
(Nielsen et al. 2007). Old forests have lower light influx,
which results in slower growing and older ramets. Most
ramets in a forest habitat are younger than 6 years old
(Flower-Ellis 1971; Nielsen et al. 2007), although 6-12
years is also common, and ramets of 34 years have also
been recorded (Flower-Ellis 1971). Shoots on single ramets
are a mix of fertile or vegetative shoots. Fertile shoots have
a lower dry weight increment than the vegetative shoots
(Tolvanen 1994), and alterations of vegetative and fertile
periods are present within single ramets (Paakonen et al.
1991). The newest part of the stem from the previous year
will be green, soft, and highly branched (Ritchie 1956).
An interesting investigation was undertaken in Belgium,
identifying 95 clones among 586 samples analyzed. Despite
of intra-population variability in the clonal diversity and in
spatial structure of the clones, no differences in clonal
diversity were detected between the three different studied
habitats. A high proportion of genetic variation existed
within populations (86%), while the variation was only 14%
between populations (Albert et al. 2004). For seedlings, the
primary rhizome does not appear until the third year at the
earliest (Sylven 1906). The present authors, however, found
that seedlings on farmland, improved by added natural peat,
grew rhizomes the second year, after seed germinating in
March and planting in July (Nestby pers. obs. 2009).
EB flowers are born singly (rarely in pairs) in the axils of
the leaves on one-year old twigs from May to June. The
twigs are sitting on aerial shots that are at least three years
old (Flower-Ellis 1971; Tirmenstein 1990; Featherstone
2002). However, sexual reproduction starts and peaks ear-
lier in rejuvenated shoots, indicating shortened ageing pro-
cess in these (Tolvanen 1994). Flower initials are developed
the year before flowering and overwinter in a dormant bud.
There are two types of buds; buds enclosing two bracts
and one single flower, and buds which bear a single lateral
shoot and a flower. These flowering shoots normally drop
together with the leaves in autumn. Some buds do not deve-
lop flowers and grow as vegetative short shoots (Flower-
Ellis 1971). With respect to floral structure, the stamens are
closed within the narrow corolla, while pistils stretch out of
the corolla. Flower diameter is approximately 4-6 mm
(Flower-Ellis 1971; Featherstone 2002). Following the pat-
tern of EB in a ten years period revealed that EB flowered
heavily every second year (Kardell and Eriksson 1990;
Selås 2000), indicating a biannual flowering pattern.
In the sweet lowbush blueberry the development is
similar. During late summer, floral bud development is ini-
tiated in the apical meristem of the upright shoot and then
proceeds down the axis of the stem in axillary buds located
in the leaf axils. However, in contradiction to the EB they
carry “clusters” or multiple inflorescences that originate
from a compound floral bud. Growers have been able to
dramatically increase yields of the sweet lowbush blueberry
by pruning the fields on alternate years. This ensures that
the fields are comprised of new shoots and thus have a high
flower density. Consequently clonal growth is likely to be
an important factor that constrains fruit and seed number
(Nuortila et al. 2002). A similar practice in EB could delay
the fruiting with several years since the generative part of
the tiller will be removed. It is a question if the shoots
sprouting from buds on the remaining long shoot initiate
flower buds the first autumn after cutting in the spring as an
effect of rejuvenation (Tolvanen 1993). Cutting may delay
sexual reproduction because of increased allocation of re-
sources to vegetative growth to replace lost tissue (Sones-
son and Callaghan 1991), and it is suggested that new EB
ramets do not produce flowers until two or three years after
cutting (Tolvanen et al. 1993a, 1993b). The effect of cutting
is under investigation by the present authors (Nestby et al.
pers. comm. 2008).
As a result of the extent of large EB clones, the rate of sel-
fing through geitonogamy (transfer of pollen between
flowers of the same genet) is significant (Albert et al. 2008).
In northern boreal forests EB is obligately insect pollinated,
and the main pollinators (bumblebee queens) make approxi-
mately 76 (Nuortila et al. 2002) to 90% (Albert et al. 2008)
of their flights within a distance of only 40 cm. The EB
have no mechanisms for avoiding self-pollination or self-
fertilization (Nuortila 2007). Flowers have been found to set
fruit equally well with self-pollen and cross-pollen (Nuor-
tila et al. 2002; Raspé et al. 2004), but cross-pollinated
flowers matured more seeds and aborted fewer seeds than
those that were self-pollinated. Approximately four times as
many seeds in cross-pollination were set at 10 m distance
when compared to self-pollinated flowers, suggesting pol-
len limitation (Jacquemart 1997) and an inbreeding depres-
sion at the seed stage (Fröborg 1996; Raspe et al. 2004;
Nourtila et al. 2006; Nuortila 2007). It appears to be a clo-
nal variation in self-fertility (Raspe et al. 2004; Nuortila
2007). Berry fresh weight is positively related with the total
weight of seeds and the number of seeds (Ranwala 2001;
Ranwala and Naylor 2004), meaning that fruit yield would
benefit from cross-pollination. Subsequently, for successful
fertilization to occur, extensive interphenotype (clone) pol-
len movement is required in fields. This indicates that for
Fig. 2 Vaccinium myrtillus. The coin is 21 mm in diameter. (Nestby
European blueberry domestication. Nestby et al.
cultivation purposes, it could be useful to expand the “win-
dows of opportunity” by in situ rhizomes or seeds or by
planting seedlings or advanced clones, to increase the repro-
ductive success and clonal diversity within pollinator flight
The most significant characteristic of blueberries is their
high content of beneficial nutrients and bioactive phytoche-
micals (Table 2). Nutritional compounds comprise carbo-
hydrates and organic acids, which mainly contribute to the
taste impression, accompanied by aroma volatiles. The
favourable berry aroma from EB shows complex phytoche-
mical patterns based on more than 100 compounds (Rohloff
et al. 2009), compared to V. corymbosum (Parliment and
Kolor 1975; Hirvi and Honkanen 1983) and V. ashei (Hor-
vat et al. 1996).
The nutraceutical quality is recognized by the abun-
dance of natural antioxidants such as proanthocyanidines
and anthocyanins (Faria et al. 2005) together with the oc-
currence of other potent flavonoids (Cho et al. 2005), other
phenols (Taruscio et al. 2004; Zadernowski et al. 2005), and
reasonable amounts of ascorbic acid (Stewart 2004). An
obvious difference is the blackish fruit flesh colour of EB
compared with the whitish colour of the American blue-
berries, including V. angustifolium, V. corymbosum L. (high-
bush), V. ashei Reade (rabbiteye) and V. myrtilloides Michx.
(sourtop lowbush). Jam made of EB has a more pronounced
blueberry-flavour and odour than that of e.g. the V. co r y m -
bosum cvs. ‘Bluecrop’ and ‘Berkely’ (Rødbotten et al.
2005). The berries of EB are characterized by 15 anthocya-
nins according to a Finnish study (Lätti et al. 2008). A sig-
nificantly lower content of the total anthocyanins was ob-
served in berries of the southern regions of Finland com-
pared with the central and northern regions, which is in
accordance with observations on latitudinal variation in
Norway (Nestby et al. 2010, unpublished). Differences in
the proportion of anthocyanins were also reported (Lätti et
al. 2008). In general, berries from EB are characterized by
higher levels of total anthocyanins, phenols, and antioxi-
dants, whereas highbush varieties show superior berry
weight and thus yield potential (Tab l e 2). The study of
berry peels and flesh with regard to the in planta distribu-
tion of bioactive phytochemicals, emphasizes the health-
beneficial properties of EB compared to highbush blue-
berries (Riihinen et al. 2008). Recently, resveratrol and
structure-relative compounds that have been reported to
show cancer-chemopreventive activities, have been des-
cribed in Vaccinium berries (Rimando et al. 2004). Ad-
ditionally, EB seed oils have been shown to serve as an
excellent source for linolenic acid, essential fatty acids,
tocopherols and carotenoids (Parry et al. 2005).
In order to fully address fruit characteristics, not only
the internal quality but also external parameters based on
morphological and physiological traits have to be con-
sidered. Blueberries from high- and lowbush cultivars show
an extended shelf-life and can be freshly stored for periods
from 4 to 6 weeks (Krupa and Tomala 2007; Echeverría et
al. 2009) when using controlled (CA) or modified atmos-
phere (MA) conditions. Due to their flesh firmness and
berry skin properties, they are fitted for long-distance trans-
port for sale on external markets far from the production
site. In contrast, berries from EB show a relatively higher
degree of vacuolization at full maturation stage, which
makes them more suitable for distribution on local markets
and fresh consumption rather than long-term storage and/or
transport. However, high levels of antimicrobial and bioac-
tive compounds in EB have high potential to suppress bac-
terial and fungal growth (Koskimaki et al. 2009). Posthar-
vest quality of blueberries is clearly reduced due to water
loss, lower firmness and shrivelling, as illustrated with V.
ashei (Schotsmans et al. 2007) and V. corymbosum (Krupa
and Tomala 2007). Whereas levels of anthocyanins and anti-
oxidant capacity decrease over time, the taste parameters of
soluble solids and titratable acidity often remain unchanged
also after several weeks of storage (Krupa and Tomala
2007). Frozen storage of blueberries as an alternative for
quality preservation of whole berries, does not significantly
change nutrient and anthocyanin composition even after 4
months (Poiana et al. 2008), when suitable freezing condi-
tions are applied.
Half cultivation
In Finland, forest owner preferences have changed from an
economy based on tree value only, to an evaluation where
also berry yields and other non-wood products have to be
taken into consideration. This has resulted in the need to
assess the effect of silvicultural treatment on berry yields
Tab le 2 Chemical composition of V. myrtillus, and other selected Vaccinium species.
Quality parameter V. myrtillus
V. myrtillus
V. corymbosum
V. angustifolium
Berry weight (mg f.w.) 457 r 81 328 r 63 1635 r 346 326 r 67
Dry matter (g/ 100 g f.w.) 15.0 r 1.6 15.2 r 3.2 16.4 r 4.1 22.1 r 13.1
Soluble solids content (Brix value in %) 10.8 r 1.6 9.8 r 1.1 12.7 r 2.1 15.4 r 1.6
pH 2.7 r 0.1 3.1 r 0.1 3.2 r 0.2 2.7 r 0.1
Titratable acidity (g/ 100 g f.w.) 1.4 r 0.2 2.4 r 1.5 1.4 r 0.6 0.9 r 0.1
Total anthocyanins (mg/ 100 g f.w.) 275 r 72 364 r 189 145 r 54 181 r 152
Total phenols (mg GAE/ 100 g f.w.) 612 r 75 472 r 164 289 r 105 546 r 255
Antioxidants FRAP (mmol/ 100 g f.w.) 5.7 r 1.2 5.3 r 2.2 2.6 r 1.1 9.8
Fructose (mg/ 100 g f.w.) 5290 r 1027 3687 r 1092 6171 r 3150 3900
Glucose (mg/ 100 g f.w.) 5348 r 1074 3380 r 988 3296 r 852 5150
Sucrose (mg/ 100 g f.w.) 578 r 270 411 r 187 180
Citric acid (mg/ 100 g f.w.) 1321 r 150 683 r 171 427 r 168
Malic acid (mg/ 100 g f.w.) 298 r 95 261 r 195
Quinic acid (mg/ 100 g f.w.) 1703 r 476 1370 46 r 73
Catechins (mg/ 100 g f.w.) 45 r 24 5.0 5.3 r
Chlorogenic acid (mg/ 100 g f.w.) 32 r 18 59 r 82
Ascorbic acid (mg/ 100 g f.w.) 3.0 r 2.5 18.6 r 24.6 9.2 r 3.8 12.1 r 6.9
Gallic acid (μg/ 100 g f.w.) 834 r 235 1760
Quercetin (μg/ 100 g f.w.) 473 r 262 2263 r 966 3824 r 2764
1Based on experimental data from Norwegian trials in 2009 (Nestby et al., unpublished data)
2Data compiled from 27 references
3Data compiled from 24 references
4Data compiled from 4 references
The European Journal of Plant Science and Biotechnology 5 (Special Issue 1), 5-16 ©2011 Global Science Books
(Kangas 1998). In Norway, this attitude has not been very
apparent. However, there has been a recent and substantial
interest in the economical value of resources other than
trees in outlying fields. Unfortunately, this has mainly foc-
used on fishing, hunting and tourism. The blueberry resour-
ces have been largely overlooked, but should be evaluated
as important. Some ecological investigations of forest man-
agement including EB, could give inputs to how this re-
source could be made more easily available for exploitation.
In addition to forest management, climate, soil conditions,
insects and diseases, browsing animals etc. have an influ-
ence on the development of the European blueberry.
Forest cutting
The EB is severely affected by clear-cutting with substantial
reductions in vegetative growth, shoot survival, ground
cover and fruit production, and an increase in shoot phenol
content. This was confirmed in a study by Atlegrim and
Sjöberg (1996) who reported a reduced ground cover and a
more patchy spatial distribution after selective cutting.
Interestingly, the vegetative investment and shoot survival
after cutting did not differ from zero cutting (Atlegrim and
Sjöberg 1996). This suggests that percentage cover of EB is
highest in established old forest. A study on different age
classes of V. myrtillus in Polish pine stands strengthen this
theory. It was shown that percentage cover- area and weight
of aerial shots were highest in 40-80 years old stands
(Kalinowski 2004). Similarly, in a pine forest of Kola, 200
years old stands of V. myrtillus had the highest production
and vitality (Maznaya 2001). This is in agreement with Kar-
dell and Eriksson (1995), who found that percentage cover
of EB increased with forest maturity, and that clear cutting
reduced percentage cover with 80%, and that the recovering
was very slow.
There is suggested a negative connection between EB
cover and irradiance; the cover was poorest at clear cutting
and greatest at intermediate irradiance, which coincided
with high crown forest stands (Parlane et al. 2006) and for-
est regularly thinned (Kardell and Eriksson 1995). However,
under Norwegian conditions the effect of clear cutting was
the opposite, and the average performance of EB decreased
with increased forest maturity and larger tree biomass. This
experience could be caused by the use of relatively small
clear cuts in Norwegian forests (Tolvanen 1994; Nielsen et
al. 2007).
Besides forest management, climate has a decisive impact
on the development of the EB. Vaccinium grow wild in the
temperate zone and stretches into the arctic climatic zone.
This environment has typical changes in seasons with rela-
tively warm summers and long days followed by colder
autumns and winters with abundant and persistent snowfall
and short days. These changes have made it necessary for
the blueberry to stop growth in late autumn, to avoid re-
sumption of growth too early in the spring, and to develop
avoidance (snow cover) and tolerance to low temperature
stress (Rowland et al. 2004). This implies that moving types
of EB from northern areas to southern areas or vice versa
probably will be unsuccessful.
In the last decades the winters have generally been mil-
der and the snow cover more unstable, at least at low alti-
tudes in Scandinavia. The importance of snow cover was
notified by Gjærevoll (1949), giving that snow cover is an
important factor which determines the altitudinal distribu-
tion of EB. In the absence of a protective layer of snow,
plants are vulnerable to cold winter temperatures and may
be killed (Hall et al. 1971), though it is moderately buffered
against frost in late winter and early spring (Tolvanen et al.
1997). In a winter five degree warmer than average in NE
Sweden during 1991-1992, V. myrtillus suffered lethal inju-
ries. It was suggested that this was connected to a conside-
rable decrease in solute content during winter (Ögren 1996).
Closer examinations including experiments with infrared
heating lamps run with or without soil warming cables,
confirmed the effects of winter warming. After a simulated
one week long extreme winter warming event in early
March, Vaccinium myrtillus had delayed bud development
by up to 3 weeks in the following spring (June) and reduced
flower production by more than 80% (Bokhorst et al. 2008).
Later this was more carefully scrutinized and tissue
water in the shoots was observed in winter compared with
spring and early summer. Soluble solids tended to decrease
from December to February under snow cover, while re-
maining constant under artificial gray and transparent cover
(roofing), indicating an enhanced demand for synthesis of
proteins, including dehydrins that protect against drought
(and freezing) under conditions without snow cover. Also
increases of metabolism in early spring were less or absent
in plants that wintered without a snow cover. This can be
explained by a delayed activation of metabolism resulting
from multiple stresses (LT, drought, light) that acted simul-
taneously on the EB plant (Tahkokorpi et al. 2007). How-
ever, it should be noticed that blueberries have dormancy
mechanisms that are present and functional during the win-
ter. The lowbush blueberry need temperatures of less than
4°C for 1600 hours for dormancy to be broken, which in
Nova Scotia typically occurs in February (Percival 2010
pers. obs.). Lack of chilling could give reduced bud burst
mistaken as winter injury. Also, nutrition would influence
winter hardiness, and Taulavuori et al. (1997) concluded
that winter hardening and glutathione status in the EB seems
to be sensitive to nitrogen fertilization, and not affected by
elevated CO2 and O3.
In a domestication strategy breeding EB-clones for win-
ter hardiness could be an objective. Little is known of in-
heritance of this character. In highbush blueberries, cold
hardiness is controlled largely by additive, multiple and
largely dominant gene effects. These genes are dehydrins
(bbdhnd1-bbdhn5) (Arora et al. 2000; Rowland et al. 2004).
Though it is not examined, probably similar mechanisms
are prevalent in the EB. However it may be anticipated that
the genes that regulate hardiness at least to some degree are
different from those found in highbush blueberries.
Another aspect worth noting is the vulnerability of
flowers to be injured by frost in late spring/early summer.
This has happened more frequently the last decades in both
lowbush and EB due to milder winters and earlier resump-
tion of growth in the spring. This frost injury can be as rate
limiting as winter injury, and changes with the developmen-
tal stage of the plant. The pattern observed in a ten year
period that EB flowered heavily every second year, could
partly be a result of this, but also due to the physiological
effects associated with biennial bearing. During registra-
tions over three years in central Finland, fruit yield varied
from zero to 130 kg ha-1, and in a Swedish 15 year study,
yields varied from 0 to 450 kg ha-1 depending on site and
year. The years with almost zero yields always had a history
of frost during bloom (Kardell and Eriksson 1990, 1995).
An examination in the Arkangels region of Russia illus-
trated that profuse flowering during fairly warm weather
with rain showers after the late spring frost (mid-June), and
enough moisture in July and August, secured high fruit
yields. These conditions typically occurred every fourth
year. In a typical good year the yield averaged from 171 to
341 kg ha-1 (Puchnina 1996).
Plant reactions that can reduce LT injury were observed
in cranberry uprights (V. macrocarpum Ait.) and fruit.
Work ma ster et al. (1999) suggested that ice nucleation of
leaves performed by ice penetration via stomata located on
the abaxial surface, and that the thick cuticle present on the
adaxial surface was an effective barrier to intrinsic nuclea-
tion. Only frozen moisture at the calyx end of the fruit in
the remnant nectary could induce fruits to freeze, most
likely through stomata. If a similar appearance is present in
EB it would be a valuable trait preserving the fruit quality
by supercooling in short periods with warm freezing tempe-
ratures in the autumn.
European blueberry domestication. Nestby et al.
Soil and nutrition
The EB is one of the dominating dwarf shrubs in forest
habitats with low nutrient availability due to its clonal
growth habit, the symbiosis with ericoid mycorrhiza (Bon-
fante-Fasolo et al 1981; Kasurinen and Holopainen 2001)
and the ability to take up organic nitrogen (Näsholm et al.
The optimal conditions for EB growth, development
and fruit yield, according to Svalestad (1983), occurred in
Norway under conditions of high humidity and minimal
shade. In addition he found that mineral nutrition increased
the fruit yield at sites where the EB was a dominating spe-
cies, especially when water was sufficient.
Similarly, in other studies, mineral nutrition was an im-
portant factor for plant productivity. In the northern Apen-
nines, total community net primary production (NPP) of
three communities was closely related to nutrient availabi-
lity. NPP of V. myrtillus peaked in the most fertile habitat,
and within this the N to P ratio in the whole plant as well as
in the leaves reflected the soil phosphate concentration with
foliar N to P ratios of less than 16 in the poorer sites. The
responses showed by other species in the same habitat sug-
gest that the response of EB is individualistic in response to
the nutrient availability, and that the growth is P-limited
(Gerdol 2005). A study of Kardell and Eriksson (1995) over
15 years showed that fertilization with 150 kg ha-1 of am-
monium nitrate the second year after establishment of trial
plots, and again before the 10th year, gave varying reactions
between field trials situated in the south and north of Swe-
den. Generally, however, fruit yield increased the first years
after implementation of fertilizer, but was reduced gradually
towards the second application of fertilizer. After the second
application a new increase in fruit yield occurred followed
again by a slow decline, but not as significant as after the
first application. The best results were achieved when ferti-
lization and thinning of trees were combined.
On the contrary to the above mentioned studies Nordin
et al. (2006) reported no effect on growth of EB after fertili-
zing a boreal forest under storey vegetation with am-
monium and nitrate in the range 12.5 to 50.0 kg N ha-1.
The uptake of nutrients in Vaccinium is facilitated by
symbioses with ericoid mycorrhiza, which provide the hosts
with access to soil-nutrient resources that would otherwise
be largely unavailable to the plant (Read 1980). Mycor-
rhizal formation is also contributing to the success of EB in
nutrient stressed environments, and Bonfante-Fasolo et al.
(1981) found that nearly all root hairs examined were mycor-
rhizal, although infection intensities varied at different
times of the year. A heavy fungal infection takes place with-
in new root formation in late summer and autumn, decrease
during winter and increase gradually again towards late
summer. The mycorrhizal colonization of Vaccinium roots
take place in the cortical cells where the fungi differentiate
typical intracellular hyphae coils, which characterize the
symbiotic association between ericaceous plants and their
symbions (Harley Smith 1983; Jeliazkova and Percival
2003a, 2003b).
In a forest several factors influence nutrient availability
of the EB. At clear-cut it is important to prevent leaching
and surface erosion of nutrients through the presence of
vegetation that retains nutrients in the ecosystem. In a
mixed forest dominated by Norway spruce (Picea abies L.)
in eastern Finland, the biomass of EB significantly de-
creased after clear cutting. However, it remained a marked
nutrient sink, and the biomass returned to initial levels soon
after clear cutting as did the nutrient contents of ground
vegetation (Palvainen et al. 2005). The fast recovery of V.
myrtillus in this study is in contradiction to the findings of
Kardell and Eriksson (1995) in a 15 years study, who found
that V. vitis-idaea had a better recovery than V. myrtillus.
The uptake of nutrients is also dependent on soil pH,
and it is shown that there is an effect on pH of type of domi-
nating tree species on blueberry land. Soil pH was higher
under birch (4.7) than under spruce (4.1), while the C/N
ratio was lower under birch (17) than under spruce (23), in
a podzol and humus type moor soil. Microbial biomass, C
and N, net N mineralization and net nitrification were all
higher under birch than under spruce, per unit organic mat-
ter (Smolander et al. 2005).
An injury of an insect may be affected by interactions of
other factors in the environment. By example the ‘Winter
moth’ larvae (Operophtera brumata, Lepidoptera) may feed
heavily on EB. At the individual level, altered food plant
quality due to repeated infection by the fungus Va ld e n si a
heterodoxa, had no effect of the larval performance in labo-
ratory experiments, but both survival to the adult stage and
adult weight were positively affected by N fertilization. In
addition exclusion of insectivorous birds increased the fre-
quency of larval damage to EB shoots, indicating higher
larval densities, and there was an indication of higher bird
predation in fertilized plants. The results suggest that top-
down effects (in this case birds) are more important for lar-
val densities than bottom-up effects (e.g. nitrogen, fungus),
and that bird predation may play an important role in popu-
lation regulation of ‘Winter moth’ in boreal forests (Streng-
bom et al. 2005). Also Agriopsis aurantiaria feed on the
leaves of EB in the lowland south in Norway. ‘Autumnal
moth’ (Epirrita autumnata) and ‘Winter moth’ are often
found together, and both may occasionally de-leaf the birch
(Betula pubescens Ehrh.) and EB totally (Fjelddalen 2009;
Wikipedia 2009b).
It is shown that the wood ant (Formica aquiloni) is
beneficial for the EB. Close to ant nests herbivore damage
to the EB was reduced and proportion of flowers succeed-
ing to berries was increased. It was therefore suggested that
distance to wood ant nests and thereby reduced predation
from ants, affect herbivore damage to the EB and its repro-
ductive success. However, vegetative growth and reproduc-
tive investment was not affected by distance to ant nests,
indicating that the EB can also compensate for losses due to
herbivore injury (Atlegrim, 2005). It should however be
noted that the red ant may feed on the blueberry flower.
This was observed in Norway. However, it seem like the in-
jury is mainly on the corolla and not on the pistil or unripe
fruit, since the berries developed normally (Nestby 2008
pers. obs.).
This suggests that when methods to improve the yield
of the EB are implemented it should be strongly considered
not to override the natural defence mechanisms, but rather
try to strengthen these or at least not weaken them.
Leaf damage caused by diseases is often observed in EB
fields. The most common is Valdensia heterodoxa. The
interaction between this parasitic fungus and its host plant
the EB, was affected by nitrogen additions over 5 years in a
boreal forest in northern Sweden. Disease incidence on
leaves increased following N addition and the effect was
stronger in large than in small plots (ranging 1 to 5000 m2),
and disease incidence was also positively correlated with
precipitation. High summer precipitation enhanced the N
effect, suggesting that precipitation may modify the effects
of N deposition on plant-parasite interactions. Parallel to
this it is observed in lowbush fields of Nova Scotia that N
increases canopy biomass, this lengthens canopy wetness
duration and greatly increases the likelihood of infection
(Percival 2010 Pers. comm.). This may complicate predic-
tions of future effects of N deposition as precipitation pat-
terns are expected to change as a result of climate change.
The results suggests that small scale fertilization experi-
ments may underestimate future large-scale effects of N-
deposition, and indicate the need for increased awareness of
the problems (Strengbom et al. 2006). N-induced changes
in the constitutive levels of soluble conjugated amines did
not seem to explain the increased parasite (Valdensia hete-
The European Journal of Plant Science and Biotechnology 5 (Special Issue 1), 5-16 ©2011 Global Science Books
rodoxa) susceptibility of the EB under N enrichment. Gene-
rally, the concentration of free diamines and insoluble con-
jugated putrescine were higher in diseased than in healthy
shoots, suggesting parasite-induced accumulation of dia-
mines. Free spermine seemed to accumulate in unfertilized,
diseased plants, but in fertilized plants this induction was
dampened, suggesting that N-induced alterations in sper-
mine metabolism may promote the spread of parasites on
the EB under N-enrichment (Witzell et al. 2005).
Mummy berry disease (Monilia vaccinii-corymbosi) is
one of the most important plant pathogenic fungi on blue-
berries in America and the first verified detection in Europe
(Austria) was in 2002. The fungus was likely introduced to
Europe years ago and is present in other blueberry plantings,
but has not been identified due to possible unfavourable
weather conditions (like low temperature) for the fungus or
confusion with similar fungi (e.g. Botrytis cinerea). It re-
mains to be seen how this disease will affect the EB, which
has some resistance to this pathogen (Gosch 2006).
Weed pressures are different in established old forests com-
pared with a clear cut sites. These pressures are also more
prevalent under birch (Betula pubescens) because of dif-
ferences in light penetration to the ground, than under Nor-
way spruce (Picea abies). Birch impose cascading effects
on both above- and below-ground communities, soil chemi-
cal and physical properties and ecosystem processes. Com-
pared with heather (Calluna vulgaris) plant species richness
decreased and the vegetation composition changed under
birch, with lower cover of grasses and EB (Mitchell et al.
When using applications of N fertilizer in EB, problems
could be encountered as a result of heather responding posi-
tively to N-application (Britton and Fisher 2008). Similar
effects were observed in clear cut areas of Norway spruce
forest during 2009, with ‘wavy hairgrass’ [Deschampsia fle-
xuosa (L.) Trin.] and ‘fireweed’ (Epilobium angustifolium)
competing strongly with the EB (Nestby 2009 pers. obs.).
Also ‘Birch’ (Betula pubescens) sprouting from stubs and
‘European Rowan’ (Sorbus aucuparia) may be a problem if
caution is not taken to prevent these to develop (Nestby
2009 pers. obs.).
Browsing animals and birds feeding on the berries
Moose (Alces alces) and deer (different species), which are
quite abundant in Norway, are feeding on EB plants (Fea-
therstone 2009). This was confirmed in an examination
undertaken in a mature Scandinavian pine (Pinus sylvestris)
forest. At sites subjected to differing natural intensities of
grazing by Cervus elaphus (red deer), it was shown that
ramet size, abundance and fruit set and invertebrate activity
on EB were negatively related to grazing intensity. Even at
low grazing intensities the performance of the plant was
affected. The effect on fruit production and invertebrate
activity indicated that red deer grazing has a negative im-
pact on the population dynamics of the plant (Tømmervik et
al. 2004; Hegland et al. 2005; Melis et al. 2006; Parlane et
al. 2006). The EB have a protective mechanism against
heavy feeding of bank vole (Clethrionomys glareolus), pos-
sibly explained by changes in plant chemistry (Selås 2006).
These browsing effects are also confirmed in studies simu-
lating herbivory (Tolvanen et al. 1993b; Tolvanen 1994).
EB foliage is highly palatable and it is also an important
food source for birds such as red grouse (Lagopus lagopus),
ptarmigan (Lagopus mutus) and black grouse (Tetrao tetrix).
The capercaillie (Tetrao urogallus) in particular depends on
EB, and it eats the stems and buds in winter, as well as the
leaves in spring and summer (Featherstone 2009). It is well
known in Scandinavia that ‘Gray thrush’ (Turdus pilaris)
and ‘Starling’ (Sturnus vulgaris) feed on the fruit (Nestby
2009 pers. obs.). Therefore, if the intention is to increase the
EB abundance, it may also be important to reduce or some-
how deter the number of browsing animals (Tømmervik et
al. 2004; Hegland et al. 2005; Melis et al. 2006; Parlane et
al. 2006).
Finally, examinations have shown that EB respond nega-
tively in growth performance and age to the disturbance and
stress on ski pistes (Rixen et al. 2004).
Cultivation on farm land
The main difference from forest fields is that the soil pH
often will have to be adjusted by adding sulphur or organic
matter of low pH (e.g. compost, natural peat). Also it may
be necessary to top-dress the soil with a layer of organic
matter at specified intervals. In addition the weed pressure
may be different from forest fields, and the protection
against light-related stress and wind given by forest trees,
will normally be lacking. However, when moving the EB
into cultivated land the possibility to adapt the cultivation
system to the plant will be more flexible. There is very little
experience with this kind of cultivation, and it is necessary
to take a broad and sequential experimental approach to
build a knowledge base.
The reproduction of EB has been widely studied due to the
use of EB for re-vegetation purposes and for its significance
in forest and heath ecology, where EB is one of the
dominating dwarf shrubs. The natural reproductive success
of EB in forest ecosystems is contributed to it being a spe-
cies with a long life span, a mixed breeding system and a
high seed dispersal potential (Flower-Ellis 1971). However,
the importance of seeds in regeneration of EB seems to
fluctuate. EB has been found to be one of two dominant
seed-bank species in five boreal forests stands in northern
Sweden (Granström 1982). On the contrary moors in Scot-
land (Ranwala and Naylor 2004) and in a closed temperate
forest of Spain (Laskurain 2004) had a complete absence of
seeds or seedlings. Interestingly, EB was one of the most
frequent shrub species in all these sites, resulting in its
clonal growth with subterranean rhizomes being essential
for spreading. Local patches of EB have been found to have
a shoot density up to 400 reproductive shoots per square
meter (Ranwala 2001). Consequently, seedling recruitment
in natural populations is possible within stands of estab-
lished conspecific adults only at “windows of opportunity”
(Erikson and Fröborg 1996). “Recruitment at windows of
opportunity” (RWO) is likely in EB since substrates suita-
ble for germination, seedling survivorship and juvenile
growth occur within the habitat space already occupied by
conspecific adults. This strategy provides that seed availa-
bility is not limiting, contributing to explain why EB each
year produce copious numbers of seeds and at the same
time is characterized by an almost total absence of seedlings.
RWO may also explain the incidence of genotypic diverse
clones represented by only one ramet that may exist within
spatially large clones (Albert et al. 2004).
The number of seeds per fruit reported in literature for
EB fruits range from an average of 25 (Flower-Ellis 1971)
to 40 (Ritchie 1956) to 71 (Ranwala and Naylor 2004). Seed
number varies greatly in response to pollination treatment
with cross-pollinated berries containing a maximum of 120
mature seeds, while self-pollinated berries containing a
maximum of 35 (Nuortila et al. 2006). Upon examining the
seeds at berry maturity, the average number of mature seeds
per berry was 66 and 12 in cross- and self-pollinated berries
Seed germination occurs over the range 15-24°C (Ran-
wala and Naylor 2004), and has been shown to occur when
stratified in the presence of light under cool (15.5°C) tem-
peratures for more than 12 weeks (Baskin et al. 2000). The
best germination has been obtained by no stratification and
16 weeks of incubation in 12 h light at 25/15°C day/night
European blueberry domestication. Nestby et al.
temperatures (Baskin et al. 2000). EB seeds are probably
conditionally dormant at maturity, as the maximum possible
germination percentage was not obtained using fresh seeds
(Baskin et al. 2000). Trials have also shown that germina-
tion is poorer in light having a high far-red ratio, as is under
a dense tree canopy (Skrindo 2005). Digestion of berries by
birds also affected EB seed germination (Honkavaara et al.
2007), while digestion by bears did not enhance germina-
tion (Skrindo 2005).
There is a considerable variation in EB fruit and hence
seed production between years. Depressed berry production
has been associated with high temperatures during flower-
bud formation in autumn, high temperatures in winter in
association with thin snow cover, frost during flowering in
spring and low or high amounts of precipitation during
berry ripening in summer (Selås 2000). Analyses of a 50-
year time series of fruit production in EB showed that the
highest production occurred in years with relatively high
levels of summer precipitation.
It is suggested that an average temperature increase will
reduce seed viability and cause negative implications for
the spread of EB in Scotland (Ranwala and Naylor 2004),
while it is likely to benefit from a warmer climate in the
subarctic (Milbau et al. 2009).
Vegetative propagation
While vegetative regeneration by rhizomes is very success-
ful in nature, vegetative propagation for making homoge-
nous plant material for cultivation and research purposes
has a low success-rate (Jaakola et al. 2001). The EB may be
propagated both in vivo and in vitro and different media
have been used. Investigating growth initiation in vitro in
EB using different concentrations of N6-isopentenyladenine
in the modified MS-medium, Jaakola et al. (2001) found
that the optimal concentration for the initiation of growth of
microshoots in vitro was 49.2 μM, compared to 24.6 and
78.4 μM. The treatment undertaken in spring, resulted in
44% growing explants after 8 weeks. Rooting of the EB
microshoots ex vitro were improved by incubating them in a
2.07 mM solution of a potassium salt of indole-3-butyric
acid (KIBA) at a concentration of 0.49 μM for 5 minutes
before planting in peat. Then 71.9% of the microshoots
rooted after 5 weeks, while 81.3% rooted in vitro in a root-
ing medium containing IBA at 0.49 μM.
Our own experiments in cooperation with University of
Natural Resources and Applied Life Science, Institute of ap-
plied Microbiology in Vienna Austria, with in vitro estab-
lishment and multiplication, have shown that clones of V.
myrtillus are successfully established and multiply with a
rate of 3-4 shoots per subculture (Laimer pers. comm. 2010).
Micropropagated plants have been shown to have a preco-
cious and higher rhizome production compared with plants
propagated from cuttings in both lingonberry (V. vitis-idaea)
(Gustavsson and Stanys 2000; Debnath 2005) and lowbush
blueberry (V. angustifolium) (Morrison et al. 2000). Rhi-
zome production in lowbush blueberry is also affected by
the N status (Smagula and Hepler 1980).
Micropropagation is reliable and efficient, especially for
the rapid introduction of new cultivars, and large-scale
liquid cultures combined with automated bioreactors could
be a tool to eliminate most manual handling in micropropa-
gation and reduce production costs significantly. Addition-
ally, molecular markers introduced in Vaccinium are power-
ful tools in the genetic identification of clonal fidelity (Deb-
nath 2009). A combination of advanced micropropagation,
including molecular markers, could be valuable in a future
establishment of new large plantings of EB on abandoned
farm land. However, experiences from the Canadian low-
bush industry have shown that the success of advanced cul-
tivars has been limited (Jamieson pers. comm. 2007.). The
reason is more political than horticultural, since the produc-
tion is advertised as exploiting the wild lowbush blueberry
growing in half cultivated fields. Anyway, in the effort to
domesticate the EB these tools should be integrated in a
scientific approach to the objective.
It would be a benefit to have a model for prediction of fruit
yield. It would provide information to decide fruit retention
and harvestability, and if this could be provided to end users
prior to harvest, it would assist in determining estimated
berry supply. In Finland Pukkula (1988) created a model
collecting data on site properties, trees and berry yields.
However, this model was dependent of a large number of
empirical data over many years and therefore, was easily
biased. Models based on expert modelling (Muhonen 1995;
Ihalainen et al. 2002, 2003, 2005) and on regression analy-
sis, to relate the expert priorities to stand characteristics,
seem to be more appropriate. However, according to Kan-
gas and Mononen (1997) expert models can only relieve the
acute need for prediction models and produce temporary
models for forest planning inexpensively and quickly. To
construct models that reliably describe the effect of trees
and the site on berry yield, one needs to gather large quan-
tities of empirical data over several years (Belonogova and
Kuchko 1979). Recently Miina et al. (2009) have construc-
ted generalized linear mixed model techniques predicting
yields as a function of site and stand characteristics, using
the permanent sample plots of the National Forest Inventory
The predictions of the expert based models correlated
logically with site and stand characteristics and were in line
with earlier models based on empiric data, and were less
laborious. According to the model, yield would be posi-
tively correlated to tree age and height and the standing vol-
ume of pine. Conversely, the yield would be negatively cor-
related to the stand (tree) basal area, the standing volume of
deciduous trees and to a Vaccinium site type or poorer
(ecological term that defines the combination of tree species
and dominating shrub species). These Finish examinations
also show that forest stands of medium fertility, but also on
rather poor mineral soil, produce the highest EB yield in
different parts of Finland. However, these findings are not
unique probably because they are subjective, and it may be
difficult to distinguish between rather poor and medium soil
sites. In addition EB thrives on poorer soil sites in northern
Finland than in southern Finland (Ihalainen et al. 2005). In
Norway it is observed that blueberry yields may be quite
high at high altitudes or in the north where birch is domi-
nant forest forming tree species, or above the tree limit
(Nestby pers. obs. 2007). This coincide the defined ‘Blue-
berry birch forest’ in Norway which is dominated by EB in
the undercover (Bendiksen et al. 2008). A long-term study
conducted in the Kirov Region of Russia showed that ave-
rage daily air temperatures and quantity of precipitation
during spring-summer had the greatest influence on EB fer-
tility. Average long-term yield of EB was highest in the
spruce EB forest (41.5 ± 4.1 g m-2) and lowest in the aspen
EB forest (21.5 ± 2.6 g m-2).
Also the fruit picking technique can affect the yield, and
picking EB fruits with a comb once or twice a year de-
creased the number of plants growing. In all picking vari-
ants including hand picking once and twice a year, mass and
area of leaves were smaller than at no picking. However,
picking twice a year with a comb increased the reproduction
value compared with no harvesting (Kalinowski 2007).
Hence, the results and observations referred to in this paper
suggest that in situations of nutrient poor environments,
blueberry growth and development is reliant upon the
mycorrhizal association for nutrient availability. However,
given the need to provide sufficient upright stem growth for
adequate floral bud induction, initiation and development,
low applications of at least nitrogen and phosphorous may
be beneficial on the poorest soils. Also, adequate amounts
of water during growth and generic development are of
The European Journal of Plant Science and Biotechnology 5 (Special Issue 1), 5-16 ©2011 Global Science Books
great importance to achieve a good fruit yield, as well as
avoidance of frost in the flower. For domestication of the
EB it will be of importance to consider the nutritional and
mycorrhizal conditions, and search optimal level of fertili-
zation and methods to strengthen the mycorrhizal associa-
tion dependent on soil conditions, to provide optimal yield,
fruit quality and plant health. A system where the ramet is
cut every second year, as practised in lowbush blueberry,
could contribute to increased fruit yield and effect the sec-
ond year pattern of EB flowering (biannual flowering), ob-
served by Kardell and Eriksson (1990), and positively con-
tribute to optimal shoot growth and flower bud production
in the sprout year. However, there are concerns that EB may
react differently, and that the regenerative shoots need more
years to develop fertile shoots.
It is indicated that medium or slightly poorer soil sites
will have the highest yield potential, and that a selective
cutting of trees is recommendable before clear cutting (at
least large clear cuttings), because that will prevent a mar-
ked reduction in EB phytomass production. Side effect is by
removing an extensive amount of forest there may be con-
cerns of removing carbon sinks and subsequently contrib.-
uting to greenhouse gas effect. There is presently no recom-
mendation on how EB and forestland can coexist for the EB
commercial production. The effects discussed above are
based on situations of stands with solely tree production,
where there are taken no steps towards weed control which
imply that all under vegetation are allowed to grow and
compete. By controlling the vegetation giving preference to
the blueberries, even richer soil sites could be suitable for
blueberry production if the soil pH was not too high (>5.2).
It is also a question how the mesomorphic character of the
EB can be handled. The EB thrives in shadowy conditions
and does not tolerate the desiccating impact of direct sun-
light (Raatikainen and Ratikainen 1983; Salo 1995). Also, it
is probably a matter of the EB undergoing excessive
amounts of photoinhibition (Percival 2010 pers. comm.).
Maybe it could be solved by cutting narrow stripes in the
forest instead of cutting a larger area leaving trees in a
spread pattern. This practice has been successfully used
with sweet lowbush blueberry production in Canada, and
should be examined in more detail with the EB. However,
in Norway growing on clear cuts show good results as well
as growing above the tree limit. This indicates that the
amount of light is not as much of a problem in EB as the
desiccative effect of sun and wind. If the water conditions
are sufficient, the EB should grow and produce well even
on large open areas. There is probably an optimal size of the
open spaces dependent e.g. on topography and altitude, also
because it is shown a reduced visit of pollinating insects
when the hedge effect of trees is lacking, and the average
temperature would also be reduced.
It is important to notice that EB is an important nutrient
sink after cutting and that the growth is markedly dependent
on P-level in the soils, which implies that the EB could
benefit from P-fertilization. However, other plant species
are effective nutrient sinks after forest clearing resulting in
the EB having to compete with these or use various technol-
ogies to minimize these grass and broadleaf weed pressures
in commercial production. An effective weed strategy also
would positively affect the fruit yield of the EB, as ob-
served in the sweet lowbush blueberry (V. angustifolium).
Other environmental factors including winter freezing, frost
during flowering, low temperatures and low rainfall during
flowering and growth and animal browsing have obviously
negative effect on fruit yields.
Aalders LE, Hall IV, Brydon AC (1979) A comparison of fruit yields of low-
bush blueberry selected clonal lines and related seedling progenies. Canadian
Journal of Plant Science 59, 857-877
Albert T, Raspé O, Jacquemart AL (2003) Clonal structure in Vaccinium myr-
tillus L. revealed by RAPD and AFLP markers. International Journal of
Plant Sciences 164, 649-655
Albert T, Raspe O, Jacquemart AL (2004) Clonal diversity and genetic struc-
ture in Vaccinium myrtillus populations from different habitats. Belgian Jour-
nal of Botany 137, 155-162
Albert T, Raspé O, Jacquemart AL (2008) Influence of clonal growth on sel-
fing rate in Vaccinium myrtillus L. Plant Biology 10, 643-649
Arora R, Rowland LJ, Lehman JS, Lim CC, Panta GR, Vorsa N (2000)
Genetic analysis of freezing tolerance in blueberry (Vaccinium section Cya-
nococcus). Theoretical and Applied Genetics 100, 690-696
Atlegrim O (2005) Indirect effects of ant predation (Hymenoptera,Formicidae)
on bilberry Vaccinium myrtillus. European Journal of Entomology 102, 175-
Atlegrim O, Sjõberg K (1996) Response of bilberry (Vaccinium myrtillus) to
clear-cutting and single-tree selection harvest in uneven-aged boreal Picea
abies forests. Forest Ecology and Management 86, 39-50
Barney DL (2008) Huckleberries and bilberry. Available online:
Baskin CC, Milberg P, Andersson L, Baskin JM (2000) Germination studies
of three dwarf shrubs (Vaccinium, Ericaceae) of Northern Hemisphere coni-
ferous forests. Canadian Journal of Botany 78, 1552-1560
Belonogova TV, Kuchko AA (1979) Plodonosenie cherniki v bereznjakah kare-
lii. Rastitelnye Resursy 15, 415-421 (in Russian)
Bokhorst S, Bjerke JW, Bowles FW, Melillo J, Callaghan TV, Phoenix GK
(2008) Impacts of extreme winter warming in the sub-Arctic, growing season
responses of dwarf shrub heathland. Global Change Biology 14, 2603-2612
Bonfante-Fasolo P, Berta P, Gianinazzi-Pearson V (1981) Ultrastructural as-
pects of endomycorrhizas in the Ericaceae. II. Host-endophyte relationships
in Vaccinium myrtilus L. New Phytologist 89, 219-224
Bendiksen E, Brandrud TE, Røsok Ø (2008) Boreale lauvskoger i Norge.
Naturverdier og udekket vernebehov. NINA rapport 367, 142-144 (in Norwe-
Britton AJ, Fisher JM (2008) Growth responses of low-alpine dwarf.shrub
heath species to nitrogen deposition and management. Environmental Pol-
lution 153, 564-573
Chen Y, Smagula JM, Litten W, Dunham S (1998) Effect of boron and cal-
cium foliar spray on pollen germination and development, fruit set, seed
development, and berry yield and quality in lowbush blueberry. Journal of
the American Society of Horticultural Science 123, 524-531
Cho MJ, Howard LR, Prior RL, Clark JR (2005) Flavonol glycosides and
antioxidant capacity of various blackberry and blueberry genotypes deter-
mined by high-performance liquid chromatography/mass spectrometry. Jour-
nal of the Science of Food and Agriculture 85, 2149-2158
Coudun C, Gegout JC (2007) Quantitative prediction of the distribution and
abundance of Vaccinium myrtillus with climatic and edaphic factors. Journal
of Vegetation Science 18, 517-524
Debnath SC (2005) A two-step procedure for adventitious shoot regeneration
from in vitro-derived lingonberry leaves: Shoot induction with TDZ and
shoot elongation using zeatin. HortScience 40, 189-192
Debnath SC (2009) Propagation and cultivation of Vaccinium species and less
known small fruits. Agronomijas Vestis 2009, 22-29
Echeverría G, Cañumir J, Serrí H (2009) Postharvest behavior of highbush
blueberry fruits cv. O’Neal cultivated with different organic fertilization
treatments. Chilean Journal of Agricultural Research 69, 391-399
Eriksson O, Fröborg H (1996) "Windows of opportunity" for recruitment in
long-lived clonal plants, experimental studies of seedling establishment in
Vaccinium shrubs. Canadian Journal of Botany 74, 1369-1374
Faria A, Oliveira J, Neves P, Gameiro P, Santos-Buelga C, de Freitas V,
Mateus N (2005) Antioxidant properties of prepared blueberry (Vaccinium
myrtillus) extracts. Journal of Agricultural and Food Chemistry 53, 6896-
Featherstone AW (2002) Species profile. Blaeberry (Vaccinium myrtillus).
Caledonian Wild! Summer 2002. Available online:
Fjelddalen J (2009) Frostmålere. Available online: Store norske leksikon,, 23.07.2010. Available online: (in
Flower-Ellis JGK (1971) Age structure and dynamics in stands of european
blueberry (Vaccinium myrtillus L.). PhD dssertation. Res. Notes 9. Royal
College For. Stockholm, 108 pp
Fröborg H (1996) Pollination and seed production in five boreal species of
Vaccinium and Andromeda (Ericaceae). Canadian Journal of Botany 74,
Gerdol R (2005) Growth performance of two deciduous Vaccinium species in
relation to nutrient status in a subalpine heath. Flora Jena 200, 168-174
Gjærevoll O (1949) Snøleievegetasjonen i Oviksfjellene. Acta Phytogeogra-
phica Suecica 25, 106 pp
Gosch C (2006) Mummy berry disease (Monilia vaccinii-corymbosi) on high-
bush blueberries in Europe. Acta Horticulturae 715, 469-472
Gough RE (1996) Blueberries- North and South. Journal of Small Fruit and
Viticulture 4, 71-106
Granström A (1982) Seed banks in five boreal forest stands originating be-
tween 1810 and 1963. Canadian Journal of Botany 60, 1821-1855
Gustavsson BA, Stanys V (2000) Field performance of 'Sanna' lingonberry
derived by micropropagation vs. stem cuttings. HortScience 35, 742-744
European blueberry domestication. Nestby et al.
Hagerup O (1928) Morphological and cytological studies of Bicornes. Dansk
Botanisk Ark 6, 1–27
Hall IV (1979) The cultivar situation in lowbush blueberry in Nova Scvotia.
Fruit Varieties Journal 33, 54-56
Hall IV, Aalders LE, Newbyrry RJ (1971) Frost injury to flowers and deve-
loping fruits of the lowbush blueberry as measured by impairment of fruit set.
Le Naturaliste Canadien 98, 1053-1057
Halvorsen BL, Holte K, Myhrstad MCW, Barikmo I, Hvattum E, Rem-
berg SF, Wold A-B, Haffner K, Baugerød H, Andersen LF, Moskaug JØ,
Jacobs DR, Blomhoff R (2002) A systematic screening of total antioxidants
in dietary plants. Journal of Nutrition 132, 461-471
Harley JL, Smith SE (1983) Mycorhizal Symbiosis, Academic Press, Inc., Lon-
don, UK, 483 pp
Hedberg I, Hedberg O (1961) Chromosome counts in British plants. Botaniske
Notater 114, 397-399
Hegland SJ, Rydgren K, Seldal T (2005) The response of Vaccinium myrtillus
to variations in grazing intensity in a Scandinavian pine forest on the island
of Svanøy. Canadian Journal of Botany 83, 1638-1644
Hirvi T, Honkanen E (1983) The aroma of some hybrids between high-bush
blueberry (Vaccinium corymbosum L.) and bog blueberry (Vaccinium uligino-
sum L.). Zeitschrift für Lebensmittel Untersuchung und Forschung 176, 346-
Hjalmarsson I (2006) Introduction of lowbush blueberry and hybrids in Swe-
den. Acta Horticulturae 715, 143-146
Honkavaara J, Siitari H, Saloranta V, Viitala J (2007) Avian seed ingestion
changes germination patterns of european blueberry, Vaccinium myrtillus.
Annales Botanici Fennici 44, 8-17
Horvat RJ, Schlotzhauer WS, Chortyk OT, Nottingham SF, Payne JA
(1996) Comparison of volatile compounds from rabbit eye blueberry (Vac-
cinium ashei) and deerberry (V. stamineum) during maturation. Journal of
Essential Oil Research 8, 645-648
Hultén E, Fries M (1986) Atlas of North European vascular plants: north of the
Tropic of Cancer I-III. - Koeltz Scientific Books, Königstein, with the kind
permission of S. Koeltz (July 2010)
Ihalainen M, Ahlo J, Kolehmainen O, Pukkula T (2002) Expert models for
bilberry and cowberry yields in Finish forests. Forest Ecology and Manage-
ment 157, 15-22
Ihalainen M, Salo K, Pukkula T (2003) Empirical prediction models for Va c -
cinium myrtillus and V. vitis-idaea berry yields in North Karelia, Finland.
Silva Fennica 37, 95-108
Ihalainen M, Pukkula T, Saastamoinen O (2005) Regional expert models for
bilberry and cowberry yields in Finland. Boreal Environment Research 10,
Ingestad T (1973) Mineral nutrient requirement of Vaccinium vitis idaea and V.
myrtillus. Physiologia Plantarum 29, 239-246
Jaakola L, Tolvanen A, Laine K, Hohtola A (2001) Effect of N6-isopentyl-
adenine concentration on growth initiation in vitro and rooting of european
blueberry and lingonberry microshoots. Plant Cell, Tissue and Organ Culture
66, 73-77
Jacquemart AL (1997) Pollen limitation in three sympatric species of Vac -
cinium (Ericaceae) in the upper Ardennes, Belgium. Plant Systematics and
Evolution 207, 159-172
Janes DE, Percival DC (2003) Trends in lowbush blueberry cultivar develop-
ment. Journal of the American Pomological Society 57, 63-69
Jeliazkova EA, Percival DC (2003a) N and P fertilizers, some growth varia-
bles, and mycorrhizae in wild blueberry (Vaccinium angustifolium Ait.). Acta
Horticulturae 626, 97-304
Jeliazkova EA, Percival DC (2003b) The seasonal tolerance of the mycor-
rhizal association of the lowbush blueberry (Vaccinium angustifolium Ait.) to
drought stress. Canadian Journal of Plant Science 83, 583-591
Kalinowski M (2004) Influence of pine stand age on chosen features of resour-
ces of Vaccinium myrtillus. Lesne Prace Badawcze 2004, 87-91
Kalinowski M (2007) The impact of bilberry Vaccinium myrtillus L. picking on
the above ground parts of plants and yield level. Lesne Prace Badawcze 2007,
Kangas J (1998) Non-wood forest goods and benefits and the choice of forest
management practices. In: Lund HG, Pajari B, Korhonen M (Eds) Sustaina-
ble Development of Non-Wood Goods and Benefits from Boreal and Cold
Temperate Forests. EFI Proceedings 23, 203-210
Kangas J, Mononen A (1997) Ekologiseen asiantuntemukseen perustuvan
numeerisen mallin tuottaminen metsäalueen biodiversiteetin arviointiin.
Metsätieteen Aikakauskirja 1997, 225-238 (in Finish)
Kangas K (2001) Wild berry utilization and markets in Finland. D.Sc. (Agr.
and For.) thesis. University of Joensuu, Faculty of Forestry, Research Notes
126, 43 pp
Kangas K, Markkanen P (2001) Factors affecting participation in wild berry
picking by rural and urban dwellers. Silva Fennica 35, 487-495
Kardell L, Eriksson L (1990) Skogsskötselmetodernas inverkan på blåbär och
lingon. Resultat av en tioårig försöksserie. SLU Rapport no. 47. Swedish
University of Agricultural Sciences, Department of Environmental Forestry
(in Swedish)
Kardell L, Eriksson L (1995) Bärprodukter och markvegetation. Effekter av
kvävegödsling och slutavverkning under en 15-årsperiod, 1977-1991. SLU
Rapport no. 60. Swedish University of Agricultural Sciences, Department of
Environmental Forestry (in Swedish)
Kasurinen A, Holopainen T (2001) Mycorrhizal colonisation of highbush
blueberry and its native relatives in central Finland. Agricultural and Food
Science in Finland 10, 113-119
Kinsman GB (1993) The history of the lowbush blueberry industry in Nova
Scotia 1950 – 1990. Blueberry Producers’ Association of Nova Scotia, March
31 1993, 154 pp
Konovalchuk VK, Konovalchuk VV (2006) The resources of wild lowbush
blueberries (Vaccinium myrtillus L., V. uligonosum) and highbush blueberry
in Ukraine. Acta Horticulturae 715, 55-59
Koskimaki JJ, Hokkanen J, Jaakola L, Suorsa, Tolonen M, Mattila S, Pirt-
tila AM, Hohtola A (2008) Flavonoid biosynthesis and degradation play a
role in early defence responses of bilberry (Vaccinium myrtillus) against bio-
tic stress. European Journal of Plant Pathology 125, 629-640
Krupa T, Tomala K (2007) Antioxidant capacity, anthocyanin content profile
in 'Bluecrop' blueberry fruit. Vegetable Crops Research Bulletin 66, 129-141
Kühn BF, Vang-Pedersen O (1991) Dyrking af blåbær. Grøn Viden Havebrug
62, 6 pp (in Danish)
Laskurain NA, Escudero A, Olano JM, Loidi J (2004) Seedling dynamics of
shrubs in a fully closed temperate forest, greater than expected. Ecography
27, 650-658
Lätti AK, Riihinen KR, Kainulainen PS (2008) Analysis of anthocyanin vari-
ation in wild populations of bilberry (Vaccinium myrtillus L) in Finland.
Journal of Agricultural Food Chemistry 56, 190-196
Lid J (1963) Norsk og svensk flora. Det norske samlaget, Oslo, Norway, 538 pp
(in Norwegian)
Maznaya EA (2001) Structure and productivity of the above-ground phytomass
of Vaccinium myrtillus L. and V. vitis-idaea L. coenopopulations in shrub-
lichen pine forests (Kola peninsula). Rastitel´nye Resursy 37, 15-22
Melis C, Buset A, Aarrestad PA, Hanssen O, Meisingset EL, Andersen R,
Moksnes A, Roskaft E (2006) Impact of red deer (Cervus elaphus) grazing
on bilberry (Vaccinium myrtillus) and composition of ground beetle (Coleop-
tera, Carabidae) assemblage. Biodiversity and Conservation 15, 2049-2059
Miina J, Hotanen JP, Salo K (2009) Modelling the abundance and temporal
variation in the production of bilberry (Vaccinium myrtillus L.) in Finnish
mineral forests. Silva Fennica 43, 577-593
Milbau A, Graae BJ, Shestova A, Nijs I (2009) Effects of a warmer climate on
seed germination in the subarctic. Annals of Botany 104, 287-296
Mitchell RJ, Campbell CD, Chapman SJ, Osler GHR, Vanbergen AJ, Ross
LC, Cameron CM, Cole L (2007) The cascading effects of birch on heather
moorland, a test for the top-down control of an ecosystem engineer. Journal
of Ecology (Oxford) 95, 540-554
Morrison SE, Smagula JM (1986) Morphology, growth, and rhizome develop-
ment of lowbush blueberry tissue culture plants, seedlings, and rooted soft-
wood cuttings. HortScience 21, 734 (Abstracts)
Muhonen T (1995) Mustikka-ja poulukkasatojen ennustaminen kasvupaikka-ja
puustotunnusten avulla-asientuntemukseen perustuva lähestymistapa. Mas-
ter’s thesis of forest planning and economies, Faculty of Forestry, University
of Joensuu, 37 pp (in Finish)
Nielsen A, Totland Ø, Ohlson M (2007) The effect of forest management ope-
rations on population performance of Vaccinium myrtillus on a landscape-
scale. Basic and Applied Ecology 8, 231-241
Nordin A, Strengbom J, Ericson L (2006) Responses to ammonium and nit-
rate additions by boreal plants and their natural enemies. Environmental Pol-
lution 141, 167-174
Nuortila C (2007) Constraints on sexual reproduction and seed set in Va c -
cinium and Campanula. Academic dissertation, university of Oulu. Acta Uni-
versitatis Ouluensis A 4 89, 60 pp. Available online:
Nourtilla C, Tuomi J, Laine K (2002) Inter-parent distance affects reproduc-
tive success in two clonal dwarf shrubs, Vaccinium myrtillus and Vaccinium
vitis-idaea (Ericaceae). Canadian Journal of Botany 80, 875-884
Nourtilla C, Tuomi J, Laine K (2006) Early acting inbreeding depression in a
clonal dwarf shrub, Vaccinium myrtillus, in a northern boreal forest. Annales
Botanici Fennici 43, 36-48
Nyström E (1932) Ett odlingsforsök med storfruktiga amerikanska blärsbuskar
i Sverige. Sveriges Pomologiska Förenings Årsskrift 1932, 180-192
Näsholm T, Ekblad A, Nordin A, Giesler R, Högberg M, Högberg P (1998)
Boreal forest plants take up organic nitrogen. Nature 392, 914-916
Ögren E (1996) Premature dehardening in Vaccinium myrtillus during a mild
winter, a cause for winter dieback. Functional Ecology 10, 724-732.
Paakonen T, Laine K, Havas P, Saari E (1988) Effects of berry production
and deblossoming on growth, carbohydrates and nitrogen compounds in Va c -
cinium myrtillus in north Finland. Acta Botanici Fennici 136, 37-42
Palvainen M, Finer L, Mannerkoski H, Piirainen S, Starr M (2005) Respon-
ses of ground vegetation species to clear-cutting in a boreal forest, above-
ground biomass and nutrient contents during the first 7 years. Ecological
Research 20, 652-660
Parlane S, Summers RW, Cowie NR, van Gardingen PR (2006) Manage-
ment proposals for bilberry in Scots pine woodland. Forest Ecology and
Management 222, 272-278
Pardo de Santayana M, Tardio J, Morales R (2005) The gathering and con-
The European Journal of Plant Science and Biotechnology 5 (Special Issue 1), 5-16 ©2011 Global Science Books
sumption of wild edible plants in the Campoo (Cantabria, Spain). Internatio-
nal Journal of Food Sciences and Nutrition 56, 529-542
Parliment TH, Kolor MG (1975) Identification of the major volatile compo-
nents of blueberry. Journal of Food Science 40, 762-763
Parry J, Su L, Luther M, Zhou KQ, Yurawecz MP, Whittaker P, Yu LL
(2005) Fatty acid composition and antioxidant properties of cold-pressed
marionberry, boysenberry, red raspberry, and blueberry seed oils. Journal of
Agricultural and Food Chemistry 53, 566-573
Poiana MA, Moigradean D, Raba D, Alda LM, Popa M (2008) The effect of
long-term frozen storage on the nutraceutical compounds, antioxidant proper-
ties and color indices of different kinds of berries. Journal of Food Agricul-
ture and Environment 8, 54-58
Puchnina LV (1996) Crop yield of fruits of Vaccinium myrtillus L. in the
Pinega national forest. Rastitel nye Resursy 32, 29-32
Pukkula T (1988) Monikäytön suunnitteluohjelmisto Monsu: ohjelmiston toi-
minta ja Käyttö. University of Joensuu, Joensu Finland, 40 pp (in Finnish,
English summary)
Raatikainen M, Ratikainen T (1983) The berry yield, picking and marketing
of Vaccinium myrtillus in the commune of Pihtipadus, northern central Fin-
land. Silva Fennica 17, 113-123 (in Finnish, English summary)
Ranwala SMW, Naylor REL (2004) Production, survival and germination of
European blueberry (Vaccinium myrtilus L.) seeds. Botanical Journal of Scot-
land 56, 55-63
Ranwala SMW (2001) Aspects of growth dynamics in European blueberry/
blueberry (Vaccinium myrtillus L.). PhD thesis, University of Aberdeen,
Aberdeen, Scotland, UK
Raspe O, Guillaume P, Jaquemart AL (2004) Inbreeding depression and
biased paternity after mixed-pollination in Vaccinium myrtillus L. (Ericaceae).
International Journal of Plant Science 165, 765-771
Read DJ (1980) The role of mycorrhizas in the nutrition of ericaceous plants
with special reference to the genus Vaccinium L. Productions Spontanees
Colloque Colmar 17-20 Juin, 191-203
Riihinen K, Jaakola L, Kärenlampi S, Hohtola A (2008) Organ-specific dis-
tribution of phenolic compounds in bilberry (Vaccinium myrtillus) and ‘north-
blue’ blueberry (Vaccinium corymbosum x V. angustifolium). Food Chemistry
110, 156-160
Rimando AM, Kalt W, Magee JB, Dewey J, Ballington JR (2004) Resvera-
trol, pterostilbene, and piceatannol in Vaccinium berries. Journal of Agricul-
tural and Food Chemistry 52, 4713-4719
Ritchie JC (1956) Biological flora of the British Isles, Vaccinium myrtillus L.
Journal of Ecology 44, 290-298
Rixen C, Casteller A, Hans Schweingruber FH, Stoeckli V (2004) Age analy-
sis helps to estimate plant performance on ski pistes. Botanica Helvetia 114,
Rohloff J, Nestby R, Nes A, Martinussen I (2009) Volatile profiles of Euro-
pean blueberry: Few major players, but complex aroma patterns. Latvian
Journal of Agronomy 12, 98-103
Rowland LJ, Panta GR, Mehra S, Permentier-Line C (2004) Molecular
genetic and physiological analysis of the cold-responsive dehydrins of blue-
berry. In: Arora R (Ed) Adaptions and Responses of Woody Plants to Environ-
mental Stresses, co-published simultaneously as Journal of Crop Improve-
ment 10, 53-76
Rødbotten M, Martinsen BK, Rosenfeldt HJ, Lea P, Haffner K (2005) Qua-
lity of highbush blueberry (Vaccinium corymbosum L.) and bilberry (Vac-
cinium myrtillus L.) jam. International Journal of Fruit Science 5, 61-71
Salo K (1995) Non-timber forest products and their utilization. In: Hytönen M
(Ed) Multiple-use forestry in the Nordic countries. The Finnish Forest Re-
search Institute 1995, 117-155
Selås V (2000) Seed production of a masting dwarf shrub, Vaccinium myrtillus,
in relation to previous reproduction and weather. Canadian Journal of Bot-
any 78, 423-429
Selås V (2006) Explaining bank vole cycles in southern Norway 1980-2004
from bilberry reports 1932-1977 and climate. Oecologia 147, 635-631
Skrindo AB, Halvorsen R (2005) Natural revegetation on forest topsoil and
subsoil along roadsides in boreal forest. Applied Vegetation Science 11, 483-
Smolander A, Loponen J, Suominen K, Kitunen V (2005) Organic matter
characteristics and C and N transformations in the humus layer of two tree
species, Betula pendula and Picea abies. Soil Biology and Biochemistry 37,
Sonesson M, Callaghan TV (1991) Strategies of survival in plants of the fen-
noscandian tundra. Arctic 44, 95-105
Stew art K (2004) Processing of cranberry, blueberry, currant, and gooseberry.
In: Barrett DM, Somogyi L, Ramaswamy H (Eds) Processing Fruits: Science
and Technology, CRC Press, Boca Raton, pp 97-112
Strengbom J, Witzell J, Nordin J, Ericson L (2005) Do multitrophic interac-
tions override N fertilization effects on Operophtera larvae? Oecologia 143,
Stri k B (2006) Blueberry production and research trends in North America.
Acta Horticulturae 715, 173-183
Svalestad SA (1983) Bærproduksjon hos blåbær (Vaccinium myrtillus), og
begrensende faktorer for denne. Thesis, ‘University of Environment and Bio-
technology’ Ås Norway, 139 pp (in Norwegian)
Sylvén N (1906) Om de svenska dikotyledonernas första förstärkningsstadium
eller utveckling från frö till blomning. Kungl. Sv. Vetenskapsakad. Hand-
lingar Bd 40 (in Swedish)
Tahkokorpi M, Taulavuori K, Laine K, Taulavuori E (2007) After-effects of
drought-related winter stress in previous and current stems of Vaccinium myr-
tillus L. Environmental and Experimental Biology 61, 85-93
Taruscio TG, Barney DL, Exon J (2004) Content and profile of flavanoid and
phenolic acid compounds in conjunction with the antioxidant capacity for a
variety of northwest Vaccinium berries. Journal of Agricultural and Food
Chemistry 52, 3169-3176
Taulavuori E, Taulavuori K, Laine K, Pakonen T, Sari E (1997) Winter har-
dening and glutathione status in the bilberry (Vaccinium myrtillus L.) in res-
ponse to trace gases (CO2 and O3) and nitrogen fertilization. Physiologia
Plantarum 101, 192-198
Tirmenstein D (1990) Vaccinium myrtillus. In: Fire Effects Information System,
U.S. Department of Agriculture, Forest Service, Rocky Mountain Research
Station, Fire Science Laboratory. Available online:
Tolvanen A (1994) Differences in recovery between a deciduous and an ever-
green ericaceous clonal dwarf shrub after simulated aboveground herbivory
and belowground damage. Canadian Journal of Botany 72, 853-859
Tolvanen A, Laine K, Pakonen P, Saari E, Havas P (1993a) Effect of habitat
and time of clipping on the recovery of the bilberry (Vaccinium myrtillus L.).
Annales Botanici Fennici 30, 15-20
Tolvanen A, Laine K, Pakonen P, Saari E, Havas P (1993b) Above ground
growth of response of the bilberry (Vaccinium myrtillus L.) to simulated her-
bivory. Flora 188, 197-202
Tolvanen A, Laine K, Pakonen P, Saari E, Havas P (1994) Response to har-
vesting intensity in a clonal dwarf shrub, the bilberry (Vaccinium myrtillus
L.). Vegetatio 110, 163-169
Tolvanen A, Laine K, Pakonen P, Saari E, Havas P (1997) Recovery of the
bilberry (Vaccinium myrtillus L.) from artificial spring and summer frost.
Plant Ecology 130, 35-39
Tømmervik H, Johansen B, Tombre I, Thannheiser D, Hogda KA, Gaare E,
Wielgolaski FE (2004) Vegetation changes in the Nordic mountain birch for-
est, the influence of grazing and climate change. Arctic- Antarctic and Alpine
Research 36, 323-332
Vander Kloet SP, Dickinson TA (1999) The taxonomy of Vaccinium section
Myrtillus (Ericaceae). Brittonia 51, 231-254
Witzell J, Kuusela T, Sarjala T (2005) Polyamine profiles of healthy and para-
site-infected Vaccinium myrtillus plants under nitrogen enrichment. Journal
of Chemical Ecology 31, 561-575
Wikipedia (2009a) Vaccinium. Available online:
Wikipedia (2009b) Epirrita_autumnata. Available online:
Workmaster BA, Palta JP, Wisniewski M (1999) Ice nucleation and propa-
gation in cranberry uprights and fruit using infrared video thermography.
Journal of the American Society of Horticultural Science 124, 619-625
Zadernowski R, Naczk M, Nesterowicz J (2005) Phenolic acid profiles in
some small berries. Journal of Agricultural and Food Chemistry 53, 2118-
... myrtillus) is characterised by purple fruit flesh. Bilberry is a native wild-growing shrub which occurs in acidic and humid forest soils in Europe, Asia and North America [2,6]. Obtaining highbush blueberry cultivars with anthocyanincontaining fruit flesh would significantly increase their value related to human health and would create the potential for the use of such fruits in the food and pharmaceutical industries. ...
... The genus Vaccinium belongs to the heather family (Ericaceae) and consists of two subgenera (Vaccinium and Oxycoccus) with about 150-450 species divided into 35 sections [6,7]. In each of the sections of high significance to breeding (Myrtillus, Cyanococcus and Oxycoccus), there are genotypes with different ploidy levels (2x, 4x and 6x). ...
... In nature, bilberry reproduces mainly vegetatively through underground rhizomes of sympodial branching, forming concentric, genetically homogeneous clusters of plants, reaching up to 15 m in diameter, with individual shrubs living up to 34 years [6]. V. myrtillus is partially self-pollinating, which leads to inbred depression [20] and could be associated with low genetic variability. ...
Full-text available
To expand the gene pool and introduce new traits to the tetraploid cultivars of Vaccinium corymbosum from wild diploid species V. myrtillus, it is necessary to double the chromosome number in diploid species in order to overcome a post zygotic crossing barrier and a strong triploid block, existing within the genus Vaccinium. Five genetically diverse bilberry genotypes were selected from 21 accessions taken from the breeding collection of the National Institute of Horticultural Research (Skierniewice, Poland) for this study. The bilberry genotypes were derived from the Polish locations of Bolimów Landscape Park, Budy Grabskie and forest complex Zwierzyniec (Łódź Province), and habitats in Norway. The selection of genotypes was made based on the analysis of amplified fragment length polymorphism (AFLP-PCR). Analysis of the Jaccard similarity indexes and the UPGMA method revealed that the examined accessions formed two main groups on the dendrogram. The first group consisted of accessions from Norway, while the second group agglomerated Polish accessions. A further two classes were distinguished in the Polish group: the first included accessions from Budy Grabskie and the second from Zwierzyniec, located ca. 9 km from Budy Grabskie. In order to obtain plant material for in vitro polyploidisation, in vitro shoot cultures of the selected accessions were initiated and multiplied. Both antimitotics used, colchicine and APM, induced tetraploids for all of the accessions. The obtained tetraploids were multiplied, rooted ex vitro and grown in a greenhouse and then in a field. The first flowering was observed in 1.5-year-old plants, either diploid or tetraploid. Diploids bloomed slightly earlier and more profusely than tetraploid plants. Compared to diploids, autotetraploids had significantly larger flowers by ca. 64% and larger pollen tetrads by ca. 35%. The germination capacity of pollen tetrads was high in tetraploids (87.8%), although slightly lower than in diploids (94.3%). After pollinating the flowers of three highbush blueberry cultivars with pollen from the bilberry tetraploid accession, J-4-4x, the plants formed fruits, some of which contained properly formed seeds. The effectiveness of interspecific crossing between V. corymbosum and tetraploid V. myrtillus, defined as the percentage of obtained seedlings in relation to the number of pollinated flowers, was highest (53.3%) in the blueberry ‘Liberty’, and lower in ‘Bluecrop’ and ‘Northland’, 14.8% and 10.0%, respectively. Before using the seedlings for further breeding, their hybridity will be confirmed by molecular markers and the phenotype will be evaluated.
... Bilberry samples from Lithuania had the lowest mean FRAP values (41.2 µmol TE/g and 40.2 µmol TE/g FW in 2019 and 2020, respectively). The FRAP results obtained in our study are close to those previously reported (53 and 57 µmol TE/g FW) in V. myrtillus fruits [50]. ...
... Bilberry samples from Lithuania had the lowest mean FRAP values (41.2 μmol TE/g and 40.2 μmol TE/g FW in 2019 and 2020, respectively). The FRAP results obtained in our study are close to those previously reported (53 and 57 μmol TE/g FW) in V. myrtillus fruits [50]. Berries showed higher antioxidant activity in the ABTS reaction system ( Figure 6). ...
Full-text available
The aim of this study was to characterize the variation in biologically active compounds, antioxidant activity and physico-chemical properties in naturally grown bilberries gathered from different sites in Northern Europe. The variability in the biologically active compounds, antioxidant capacity and physico-chemical properties, as well as the development of tools for the authenticity and quality control of wild bilberries (V. myrtillus L.) in different geographical locations was evaluated. The berries of bilberries were handpicked during the summers of 2019 and 2020 during the time periods when they are typically harvested for commercial purposes in Northern Europe (Norway (NOR), Finland (FIN), Latvia (LVA) and Lithuania (LTU)). Berries from locations in NOR were distinguished by their higher mean TPC (791 mg/100 g FW, average), whereas the mean TPC of samples from the most southern country, LTU, was the lowest (587 mg/100 g FW). The TPC of bilberries ranged from 452 to 902 mg/100 g FW. The TAC values of investigated bilberry samples varied from 233 to 476 mg/100 g FW. A high positive correlation was found between TPC and antioxidant activity of the bilberry samples (R = 0.88 and 0.91 (FRAP and ABTS assays, respectively)), whereas the correlation between TAC and antioxidant activity was lower (R = 0.65 and 0.60). There were variations in the TPC and TAC values of investigated berries, suggesting that genotype also affects the TPC and TAC in berries. In 2020, the pH values and TSS contents of berries were significantly lower than in 2019. To the best of our knowledge, this is the first comprehensive reported evaluation of the biologically active compounds in wild bilberries from different Northern European countries using one laboratory-validated method.
... In some Nordic countries with rich berry resources, such as Finland, Norway, and Sweden, the forest economical preference has progressively changed from the style based on tree value only to the model considering berry yields as well. 10,11 Wild bilberry (Vaccinium myrtillus, later termed pigmented or blue bilberry (BB)), also known as the European blueberry, is among the most economically valuable wild berries in Northern Europe and is well-known for its richness of antioxidants with health promoting effects, in particular of anthocyanins. 12 In comparison with several of its closely related small berry species, for example blueberry (V. ...
... Along with the development of high-value-added bilberry products, like BB wine, breeders have begun to cultivate BB on arable land through a series of practices, such as fertilization, irrigation, weed control, and modulation of soil pH, to extend the output of this berry. 10,15,16 In comparison with the variety of pigmented bilberry, the anthocyanin-free white bilberry (WB) is a natural variation with significantly lower expression of structural genes, particularly chalcone synthase (CHS), flavanone 3-hydroxylase (F3H), dihydroflavonol 4-reductase (DFR), anthocyanidin synthase (ANS), and flavonoid 3-O-glycosyltransferase (FGT) (Figure S1), and the strongly down-regulated expression of VuMYBPA1 and VuMYBC2 transcription factors. 8,17 Beside white bilberry, mutants of white bog bilberry and white and green currants (Ribes spp.) have also received significant scrutiny from researchers. ...
Full-text available
The nonanthocyanin phenolic compounds in juice and wine produced from fruits of white bilberry, a nonpigmented mutant of Vaccinium myrtillus, and blue bilberry (pigmented variety) were analyzed using liquid chromatography with a diode array detector (LC-DAD) and LC-DAD−electrospray ionization-quadrapole/time of flight hybrid mass spectrometry (ESI-QTOF-MS). On the basis of elution order, UV−vis spectra, accurate mass data, and fragmentation pattern and standards, 42 compounds including 22 phenolic acids, 15 flavonols, and 5 flavan-3-ols, were identified in juices and wines prepared from the two bilberry varieties. The levels of most individual nonanthocyanin phenolic compounds in white bilberry products were significantly lower than those in pigmented ones. In bilberry juices, phenolic acids were the most predominant, accounting for approximately 80% of total phenolic content, with p-coumaroyl monotropeins and caffeic acid hexoside being the major phenolic acids. After fermentation, the total contents of phenolic acids, flavonols, and nonanthocyanin phenolic compounds significantly increased, while the content of total flavan-3-ols decreased significantly. p-Coumaroyl monotropeins still dominated in the wine products, while caffeic acid content showed dramatic elevation with the significant drop of caffeic acid hexoside.
... The Vaccinium genus consists of woody perennial shrubs belonging to the Ericaceae family and contains, among other species, the Vaccinium myrtillus L. This shrub varies in size from approximately 5 to 90 cm; the flowers are bellshaped, white, pale pink, or red, sometimes tinged greenish, producing a blue-colored berry, known as blueberry [1]. This species is well known, and its fruits are highly consumed, not only for their organoleptic characteristics but also for their medicinal properties, given their richness in bioactive molecules such as phenolic compounds [2]. ...
Full-text available
Purpose The interest in using bioresidues produced by the agri-food industry is increasingly intrinsic to the world economy. As such, researchers started to look for new ways to enhance its use, developing innovations to transform these residues into high-value raw materials with industrial application, promoting the circular economy concept. Methods Therefore, pruning residues from the production of Vaccinium myrtillus L. could be an interesting exploitation field, given its position as a superfood due to its well-known antioxidant properties. The antioxidant, antibacterial, anti-inflammatory, anti-tyrosinase, and cytotoxicity potentials of aqueous and hydroethanolic extracts of V. myrtillus aerial parts were evaluated. Results From the obtained results, ultrasound-assisted and maceration extracts were found to be as effective as, if not more effective than, conventional antibiotics against methicillin-resistant Staphylococcus aureus. Aside from antibacterial activity, the extracts also showed antioxidant and anti-tyrosinase effects, which were found to be favourably related to the level of caffeoylquinic acid derivatives. Conclusion The obtained results highlight the bioactive potential and the importance of exploiting this bio residue as a novel candidate for industrial application, taking advantage of their biological properties. Graphical Abstract
Full-text available
Cuticle is the first layer protecting plants against external biotic and abiotic factors and is responsive to climatic factors as well as determined by genetic adaptations. In this study, the chemical composition of bilberry fruit cuticular wax was investigated through a latitudinal gradient from Latvia (56°N 24°E) through Finland (65°N 25°E) to northern Norway (69°N 18°E) in two seasons 2018 and 2019. Changes in the major cuticular wax compounds, including triterpenoids, fatty acids, alkanes, aldehydes, ketones, and primary alcohols, were detected by GC-MS analysis. Generally, a decreasing trend in the proportion of triterpenoids from southern to northern latitudes, accompanied with an increase in proportion of fatty acids, aldehydes, and alkanes, in bilberry fruit cuticular wax was observed. A correlation analysis between climatic factors with proportion of wax compounds indicated that temperature was the main factor affecting the cuticular wax composition in bilberries. A controlled phytotron experiment with southern and northern bilberry ecotypes confirmed the major effect of temperature on bilberry fruit cuticular wax load and composition. Elevated temperature increased wax load most in berries of northern ecotypes. The level of triterpenoids was higher, while levels of fatty acids and alkanes were lower, in wax of bilberry fruits ripened at 18°C compared to 12°C in both northern and southern ecotypes. Based on our results, it can be postulated that the predicted increase in temperature due to climate change leads to alterations in fruit cuticular wax load and composition. In northern ecotypes, the alterations were especially evident.
Development of primary spruce forests is driven by a series of disturbances, which also have an important influence on the understorey vegetation and its diversity. Early post-disturbance processes have been intensively studied, however, very little is known about the long-term effects of disturbances on the understorey. We quantified disturbance history using dendrochronological methods to investigate its impact on vascular plant diversity and understorey species composition. We sampled 141 plots randomly assigned throughout primary stands located in the zone of natural montane acidophilous forests dominated by Picea abies (L.) Karst. in the Western Carpathians. Dendrochronological, dendrometric, and environmental parameters were related to understorey properties using ordination methods and a Bayesian approach using multilevel linear models (GLMM). Time since the last disturbance (23–260 years ago; mostly windstorms and bark beetle outbreaks) had a significant effect on understorey species composition of the current communities, and it also interacted with disturbance severity to influence species diversity. The effect of disturbances on the understorey was largely mediated by the alteration of stand structure (age, DBH, canopy openness), Vaccinium myrtillus L. cover, and topsoil chemical properties. A period of severe disturbances between 1860 and 1890 resulted in a legacy of our current, relatively homogeneous spruce stands with less diverse sciophilous understorey dominated by V. myrtillus, which is in contrast to heterogeneous stands (in terms of age and spatial structure) driven by small-scale, lower-severity disturbances, which led to an understorey enriched by species with higher demands on light and topsoil quality (higher K concentration and lower C/N ratio). All developmental pathways following disturbances create a unique complex of spatiotemporal understorey variability in the montane spruce forests. Therefore, to preserve their full diversity, disturbances of all severities and sizes should be accepted as natural drivers, both in the field of nature conservation and close-to-nature forestry efforts.
Full-text available
Bilberry (Vaccinium myrtillus L.) fruits are an important part of local diets in many countries and are used as a medicinal herb to treat various disorders. Extracts from fruits are often a part of eye health-promoting supplements, whereas extracts from leaves are advertised for type 2 diabetes mellitus and glycemic control. This review provides an overview of the current knowledge of the phytochemical contents of bilberry fruits and leaves and their bioactivities, critically summarizes origins of the health claims and the outcome of clinical trials, with special attention towards those published in the past 10 years. Overall, the three most referenced indications, which are type 2 diabetes mellitus, vision disorders and circulatory diseases, all include contradictory results with no clear conclusion as to the benefits and recommended dosages. Moreover, the indications for vision disorders and diabetes originate from unproven or false claims that have been repeated in research since the 20th century without consistent fact-checking. Beneficial clinical results have been attested for the treatment of dyslipidemia and chronic inflammatory disorders when applied as dietary supplementation of fresh bilberries or as anthocyanin-rich bilberry fruit extracts. However, there is a general lack of double-blinded controlled research with larger sample sizes.
In this paper the results of spectroscopic studies and nonlinear optical properties characterization of blackcurrant and bilberry natural tinctures as well as their solutions and thin films with deoxyribonucleic acid (DNA) are reported. The spectroscopic properties were characterized by UV–Vis and fluorescence analyses. The third order nonlinear optical properties (NLO) of DNA bio tinctures were determined by the third harmonic generation technique at 1064.2 nm. Their optical damage thresholds have been also calculated.
Natural extracts are getting more attention due to the fast degradability rate and low toxicity. In this paper the obtained results for the complex based on deoxyribonucleic acid biopolymer with bilberry (BBE) natural extract are presented and discussed. The interaction between the two materials (DNA-BBE complex) was evaluated by measuring the linear optical properties using UV–Vis and fluorescence spectral methods. Furthermore, the DNA-BBE complex was used for non-linear optical (NLO) properties investigations of the obtained thin films on glass substrate by spin-coating technique. Also, the optical damage threshold of the studied DNA-BBE complexes was measured in order to check their potentiality and interest for application in photonics.
The winter moth, Operophtera brumata (L.) is an invasive forest and agricultural pest in North America that causes severe defoliation to a wide range of host species. This study examines the differential larval densities, development, and survival on seven host species in midcoast Maine: red oak (Quercus rubra L., Fagales: Fagaceae), apple (Malus domestica L., Rosales: Rosaceae) and crab apple (Malus sp. L., Rosales: Rosaceae), red maple (Acer rubrum L., Sapindales: Sapindaceae), pin cherry (Prunus pensylvanica L., Rosales: Rosaceae), white birch (Betula papyrifera L., Fagales: Betulaceae), wild lowbush blueberry (Vaccinium angustiflolium L., Ericales: Ericaceae), and highbush blueberry (Vaccinium corymbosum L., Ericales: Ericaceae). We also explore the degree of synchrony between selected host plants and larval hatch and its effect on survival. We found that densities, development, and survival were significantly greater on red oak (Quercus rubra) and apple (Malus sp.) than on all other target species and were lowest on pin cherry (Prunus pennsylvanica). We found low larval densities in open, wild lowbush blueberry fields; however, larvae successfully fed and developed on wild lowbush blueberry in a laboratory setting. This suggests that winter moth is a potential pest to wild lowbush blueberry in Maine if the outbreak expands to include areas with wild lowbush blueberry production.
Full-text available
Growth of bilberry was investigated in two boreal forest types (moderately dry pine heath forest, EM, and mesic spruce heath forest, HM) after being clipped at ground level in July, August of September 1985. Treatments simulated herbivory damage caused by small rodents. Regrowth during the same season occurred after clipping in July, but survival of these ramets was low, especially in HM forest. The two later clipping times prevented current-year regrowth, but allowed new shoots to emerge in the follwing years. Calculated as a percentage of the original value, the biomass that had accumulated by 1989 was greater in the HM forest than in the EM forest. In the EM forest, the density and the biomass reached the control level in the plots clipped in July, growth decreasing as the time of clipping became later. Ramets were smaller in the EM forest, but the percentage of leaf biomass to total biomass was somewhat greater than in the HM forest. Results suggested that the ability of the bilberry to recover completely from severe herbivory between the years of peak rodent populations depends both on the timing of the damage and on the growing site. -from Authors
Conference Paper
Full-text available
In the last 10 years, the area planted to all blueberries in North American increased 30% to 96,869 ha. During this period, lowbush blueberry area increased 33% and highbush 22%. In the USA, the area planted to "highbush" (northern and southern highbush and rabbiteye) blueberries increased from 19,758 to 22,393 ha from 1992 to 2003, a 13% increase. In 2003, the Midwest region of the USA accounted for 36% of the area of highbush blueberries planted. The South, New England, and Western regions accounted for 28%, 17%, and 14% of the planted area in 2003, respectively. Specific states in the USA that had considerable growth from 1992 to 2003 were California, Florida, Mississippi, North Carolina, Oregon, and Washington. In Canada, the area planted to highbush blueberries increased 102% to 4397 ha. Commercial blueberry plantings in Mexico were estimated at 28 ha in 2003. In the USA, total lowbush area increased 6% since 1992 with Maine accounting for 97% of the area planted. In Canada, lowbush area increased 57% since 1992 with 37% and 34% of the area planted in Quebec and Nova Scotia, respectively. The blueberry industry is still projected to grow considerably in the next 5 to 10 years. Highbush blueberries in the USA are expected to increase 16% and 35% in the next 5 or 10 years, respectively. In Canada, planted area of highbush blueberries is expected to increase by 22% in 5 years and 25% in 10 years. If projections are correct, planted area in Mexico will increase by almost 30 fold in 10 years. The managed area of lowbush blueberries is expected to increase by 10% to 20% in the next 5 to 10 years. Data on typical yields, types of cultivars grown, markets, proportion of machine harvest, major production problems, changes in production practices and research areas are presented.
Full-text available
The effects of the Individual Quick Freezing (IQF) process and frozen storage at -18° C up to 10 months, on the nutraceutical compounds, antioxidant properties and color indices of various berries (blueberry, red raspberry and blackberry) have been evaluated. Samples were extracted and analyzed for their total phenolics content, total monomeric anthocyanins, vitamin C, antioxidant activity and color indices. Total anthocyanins and color indices were evaluated by using pH-differential method, total phenolics content was measured using Folin-Ciocalteu procedure, vitamin C content using 2,6-dichlorophenolindophenol method and antioxidant activity using ferric-reducing antioxidant power (FRAP) assay. Blueberry contains the highest amounts of polyphenols, anthocyanins and antioxidant activity among the berries studied. The highest content of vitamin C was found in fresh raspberry. After freezing, no significant difference was observed for investigated nutraceuticals and color of berries, because the IQF is a rapid and non-destroying preservation method. Results showed that the frozen storage up to 4 months did not significantly affect the bioactive compounds and color indices of berries. The degradation of these characteristics was not recorded more than 23% during six months of storage. After 10 months, the content of polyphenols decreased up to 28-47% of the initial value; the total anthocyanins was found in proportion of 80-91%, and the ascorbic acid content was kept at 62-76%. After 10 months of storage the smallest loss of antioxidant activity was recorded for blueberries (approximately 23%) and the biggest loss for raspberries (approximately 37%). The results showed a positive correlation between antioxidant capacity and polyphenols, vitamin C and anthocyanins content. The correlation coefficient between FRAP and the total phenolics was higher than the correlation coefficient between FRAP and total anthocyanins or FRAP and vitamin C for all investigated berries.
Full-text available
In extreme arctic environments, selection forces driving evolution are mainly of the phsyical environment and plant interactions are positive. Elsewhere, biotic factors, such as herbivory, are important and plant interactions become negative through competition. Low winter temperatures rarely affect arctic plants, but snow depth and duration influence species distributions. Deep and persistent snow deforms plants and limits the period of resource acquisition. Cryptogams are common in such snow beds. Little or no snow cover exposes plants to abrasion by wind-blown particles and desiccation. In such fell-field sites, deciduous species and xerophytes, such as evergreen cushion plants, are common. Arctic summers are short and developmental processes are extended beyond one growing season, with perennials predominating. Cushion plants efficiently increase their temperatures above ambient, while evergreen and deciduous ericaceous dwarf shrubs exist and have complementary strategies for intercepting radiation in a low canopy. Tundra soils are generally infertile and may be disturbed by freeze/thaw cycles. Nutrients are conserved by recycling within shoots and between ramets within clones. Vegetative proliferation enhances the survival of young ramets, while physiological integration between ramets enables young ramets to forage across patchy environments. Periodic infestations of moth caterpillars defoliate large areas of mountain birch Betula pubescens tortuosa and stimulate increases in populations of their predators. Periodic population peaks of small rodents graze or kill much vegetation and they may moderate the dynamic structure of plant communities, as the plant species have different abilities to regenerate. -from Authors
In 1977, the total yield of bilberry in Pihtipudas was 2.4 million kg or 22 kg per hectare of land area. Only 2 per cent of the total yield of bilberry was picked 72 per cent of the bilberries picked were used by the families who picked them, 24 per cent were sold and 4 per cent were used for other purposes. -from English summary
Either all annual branches except the oldest stem were removed (R1), or half of the branches were removed (R2). Above-ground growth and fecundity were monitored for three growth seasons. Survival was >80%. The above-ground vegetative growth was vigorous, being 1.3-1.4 or 1.7-fold for R1 and R2 ramets, respectively, compared with control ramets. Increase in biomass in R1 ramets was a conseqence of the larger sizes of the annual branches and leaves, whereas in R2 ramets both the number and the size of the branches and leaves increased. Flower production decreased during the following growth seasons in R2 ramets, and fewer flowers developed into berries. Abundant sexual reproduction would apparently have been too costly for the stressed plants. R1 rejuvenated the ramets to a vegetative state; these started producing berries only in the third season following damage. -from Authors
Up to 39 tonnes/ha were obtained in three-harvest cumulative yields of superior lowbush blueberry materials planted in 1972 and harvested in 1974, 1976 and in 1978. Plants from four select clones yielded an average of 50% more fruit than did their four respective seedling progenies. At a spacing of 20 000 plants/ha, 70% survived through the third fruiting season.