Sea Buckthorn (Hippophae rhamnoides L.): A Multipurpose Plant

Abstract

Sea buckthorn (Hippophae rhamnoides L.) is a multipurpose, hardy, deciduous shrub, an ideal plant for soil erosion control, land reclamation, wildlife habitat enhancement, and farmstead protection. It has high nutritional and medicinal values for humans. The majority of sea buckthorn research has been conducted in Asia and Europe. It is a promising new crop for North America, and recently it has attracted considerable attention by researchers, producers, and industry.

Full text

HortTechnology ● Oct./Dec. 1996 6(4)370
Review
Sea Buckthorn (
Hippophae
rhamnoides
L.): A Multipurpose Plant
Thomas S.C. Li1 and W.R. Schroeder2
AA
AA
ADDITIONALDDITIONAL
DDITIONALDDITIONAL
DDITIONAL
INDEXINDEX
INDEXINDEX
INDEX
WORDSWORDS
WORDSWORDS
WORDS. Soil erosion, land reclamation, farmstead protection,
nutritional and medicinal values
SS
SS
SUMMARYUMMARY
UMMARYUMMARY
UMMARY..
..
. Sea buckthorn (Hippophae rhamnoides L.) is a multipurpose, hardy,
deciduous shrub, an ideal plant for soil erosion control, land reclamation,
wildlife habitat enhancement, and farmstead protection. It has high nutritional
and medicinal values for humans. The majority of sea buckthorn research has
been conducted in Asia and Europe. It is a promising new crop for North
America, and recently it has attracted considerable attention by researchers,
producers, and industry.
Fiedler, 1981; Kondrashov and
Sokolova, 1990); however, most natu-
ral populations grow in areas receiving
400 to 600 mm of annual precipita-
tion. Myakushko et al. (1986) recom-
mended that sea buckthorn not be
grown on dry soils, and Lu (1992)
noted the need for irrigation in regions
receiving <400 mm of rainfall per year.
Some species or subspecies of sea buck-
thorn can endure inundation but can-
not be grown on heavy, waterlogged
soils (Myakushko et al., 1986), al-
though they take up water rapidly
(Heinze and Fiedler, 1981). Sea buck-
thorn develops an extensive root sys-
tem rapidly and is therefore an ideal
plant for preventing soil erosion
(Cireasa, 1986; Yao and Tigerstedt,
1994). It also has been used in land
reclamation (Egyed-Balint and Terpo,
1983; Kluczynski, 1979; Schroeder
and Yao, 1995) for its ability to fix
nitrogen and conserve other essential
nutrients (Akkermans et al., 1983;
Andreeva et al., 1982; Dobritsa and
Novik, 1992)
Sea buckthorn was imported origi-
nally into Canada from Russia to the
Morden Research Station, Agriculture
and Agri-Food Canada, Morden,
Manitoba, in 1938 (Davidson et al.,
1994). Plantings were limited to orna-
mental landscapes, except in the prov-
inces of Saskatchewan and Manitoba.
The Shelterbelt Centre of the Prairie
Farm Rehabilitation Administration
(PFRA) of Agriculture and Agri-Food
Canada (Indian Head, Saskatchewan)
have been growing sea buckthorn for
30 years. It is one of the hardiest and
most adaptable woody plants used in
prairie conservation programs
(Schroeder, 1988). More than one
million seedlings have been distrib-
uted, and more than 250,000 mature
fruit-producing plants grow on the
prairies for enhancement of wildlife
habitat, farmstead protection (Pear-
son and Rogers, 1962), erosion con-
trol (Cireasa, 1986), and marginal land
reclamation (Balint et al., 1989;
Kluchzynski, 1989; Schroeder, 1990).
Sea buckthorn can be used for
many purposes (Fig. 3) and, thus, has
considerable economic potential. A
natural sea buckthorn habitat can yield
750 to 1500 kg·ha–1 of berries (Lu,
1992), shelterbelt plantings 4 to 5
t·ha–1 (Schroeder and Yao, 1995), and
orchard plantings, up to 10 t·ha–1 (C.
McLoughlin, personal communica-
tion). The vitamin C and E contents
out temperate zones between 27° and
69° N latitude and 7° W and 122° E
longitude (Pan et al., 1989; Rousi,
1971) including China, Mongolia,
Russia, Great Britain, France, Den-
mark, Netherlands, Germany, Poland,
Finland, Sweden, and Norway (Fig. 2)
(Wahlberg and Jeppsson, 1990; Yao
and Tigerstedt, 1995). During the last
decade, it has attracted considerable
attention from researchers around the
world, and recently in North America,
mainly for its nutritional and medici-
nal value.
Sea buckthorn can be cultivated,
but fails to set fruit, at an altitude of
3900 m (Rousi, 1971). In Russia, large,
native populations grow at altitudes of
1200 to 2000 m above sea level (Eliseev
and Fefelov, 1977). It can withstand
temperatures from –43 to 40 °C (Lu,
1992). Sea buckthorn is considered to
be drought resistant (Heinze and
1Agriculture and Agri-Food Canada, Research Centre,
Summerland, B.C., Canada VOH 1ZO.
2Agriculture and Agri-Food Canada, P.F.R.A.,
Shelterbelt Centre, Indian Head, Saskatchewan, Canada
SOG 2K2.
Contribution no. 971.
The cost of publishing this paper was defrayed in part by
the payment of page charges. Under postal regulations,
this paper therefore must be hereby marked advertise-
ment solely to indicate this fact.
Sea buckthorn (Hippophae
rhamnoides) is a hardy, de-
ciduous shrub with yellow or
orange berries (Fig. 1) (Bailey and
Bailey, 1978), which has been used for
centuries in Europe and Asia. In an-
cient Greece, leaves of sea buckthorn
added to horse fodder was well re-
puted to result in weight gain and
shiny hair; thus, the Latin name ‘Hip-
pophae’ meaning shining horse (Lu,
1992). Sea buckthorn occurs as a na-
tive plant distributed widely through-

HortTechnology ● Oct./Dec. 1996 6(4) 371
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are as high as 600 and 160 mg/100 g
of fruit, respectively (Bernath and
Foldesi, 1992). Its pulp and seeds con-
tain essential oil important for its me-
dicinal value, such as superoxide
dismutase activity in mice, and it has
enhanced the activity of NK cells in
tumor-bearing mice (Chen, 1991; Dai
et al., 1987; Degtyareva et al., 1991).
Taxonomy
Hippophae belongs to the family
Elaeagnaceae. Arne Rousi (1971) clas-
sified Hippophae (2n = 24) into three
species based on morphological varia-
tions: H. rhamnoides L., H. salicifolia
D. Don, and H. tibetana Schlecht.
Hippophae rhamnoides was divided
further into nine subspecies: carpatica,
caucasica, gyantsensis, mongolica, sin-
ensis, turkestanica, yunnanensis,
rhamnoides, and fluviatilis Rousi. Liu
and He (1978) described a fourth
species, H. neurocarpa Liu & He,
found on the Qinghai-Xizang plateau
of China. These classifications were
accepted generally by Lian (1988),
but taxonomists still disagree on the
precise classification of the genus.
Natural distribution
Sea buckthorn is native to Europe
and Asia (Fig. 2). The total area of sea
buckthorn in China, Mongolia, and
Russia is about 810,000 ha of natural
stands and 300,000 to 500,000 ha
planted (Sun, 1995). Natural sea buck-
thorn stands are also widespread in Eu-
rope—on river banks and coastal dunes
along the Baltic Coast of Finland, Po-
land, and Germany (Biswas and Biswas,
1980; Kluczynski, 1989; Rousi, 1971)
and on the western coast and along the
Gulf of Bothnia in Sweden. In Asia, sea
buckthorn is distributed widely through-
out the Himalayan regions including
India, Nepal, and Bhutan and in the
northern parts of Pakistan and Afghani-
stan (Lu, 1992).
Description
Sea buckthorn is a deciduous, dio-
ecious shrub, usually spinescent, reach-
ing 2 to 4 m in height. It has brown or
black rough bark and a thick grayish-
green crown. Leaves are alternate, nar-
row, and lanceolate with a silver-gray
color on the upper side (Synge, 1974).
This calcicolous species tolerates low
temperatures, high soil pH of 8.0, and
salt spray (Bond, 1983). The plant’s
extensive root system is capable of
holding the soil on fragile slopes. Sea
buckthorn can be planted in marginal
soils due to its symbiotic association
with nitrogen-fixing actinomycetes
(Akkermans et al., 1983; Dobritsa and
Novik, 1992). Roots of sea buckthorn
also are able to transform insoluble
organic and mineral matter in the soil
into more soluble states (Lu, 1992).
The plant rapidly spreads by rhizoma-
tous roots, and will quickly colonize
adjacent areas.
The sex of seedlings cannot be
ascertained until they start to flower
(Synge, 1974). Flower buds are formed
mostly on 3-year-old wood, differenti-
ated during the previous growing sea-
son (Bernath and Foldesi, 1992). The
male inflorescence consists of four to
six apetalous flowers. Pollen is released
in large quantities when the air tem-
perature reaches 6 to 10 °C. The fe-
male inflorescence usually consists of
one single apetalous flower with one
ovary and one ovule. The plant de-
pends entirely on the wind for pollina-
tion: neither the male nor the female
flowers have nectaries and do not at-
tract insects.
Cultural management
Sea buckthorn normally is trans-
planted or directly seeded in the spring.
Best growth occurs in deep, well
drained, sandy loam soil with ample
organic matter (Wolf and Wegert,
1993). In arid or semiarid areas, water
must be supplied for establishment.
Information in the literature re-
garding the cultivation of sea buck-
thorn is limited. In Saskatchewan,
Canada, seedlings planted in
shelterbelts are often under stress due
to lack of proper management. For
commercial production in orchard-like
plantations, cultural management is
clearly important. Good growing con-
ditions produce higher yield and good-
quality fruit (Walhberg and Jeppsson,
1990; Wolfe and Wegert, 1993). Crop
management of sea buckthorn should
include fertilization and cultural prac-
tices such as spacing, pruning, irriga-
tion, and weed control.
SOIL TEXTURE AND PH. In its
natural environment, sea buckthorn
plants are found on slopes, riverbanks,
and seashores. Soil acidity and alkalin-
ity, except at extreme levels ,are not
limiting factors. In China, plants have
been found in soils ranging from pH
5.5 to 8.3, although Lu (1992) re-
ported that sea buckthorn thrives best
at pH 6 to 7. Wolf and Wegert (1993)
reported actinomycetes, which have a
low tolerance for acid soil, living in
symbiosis with sea buckthorn, which
has a favorable range of pH 5.4 to 7.0.
Sea buckthorn is salt tolerant. An
unconfirmed report indicated that
0.15% NaCl added to the growing
media increased the growth of some
varieties of sea buckthorn in the labo-
ratory (Lu, 1992). Furthermore, soak-
ing seeds for 24 h in 0.15% NaCl
before sowing increased the number
of vigorous seedlings.
PRECIPITATION AND SOIL MOIS-
TURE. Most natural populations of sea
buckthorn grow in areas receiving an-
Fig. 1. Sea buckthorn berries.

HortTechnology ● Oct./Dec. 1996 6(4)372
nual precipitation of 400 to 600 mm.
For economic reasons, sea buckthorn
orchards should be restricted to areas
receiving a minimum of 400 mm of
annual precipitation. Sea buckthorn is
sensitive to severe soil moisture defi-
cits, especially in spring when plants
are flowering and young fruit are be-
ginning to develop. Under extended
drought situations (i.e., >30 centibars),
young fruit may dehydrate or abscise
(Lu, 1992). Buglova (1978) reported
that weather conditions, especially pre-
cipitation, could affect fruit weight. In
Belarus Academy of Science, Russia,
Garanovich (1995) reported that irri-
gation was necessary during dry grow-
ing seasons. In an unconfirmed report,
Lu (1992) indicated that the mini-
mum soil moisture levels needed to
cultivate sea buckthorn in medium
clay loam, heavy clay, slightly sandy
soil, and sandy loam were 70%, 80%,
60%, and 65% to 70%, respectively. Li
(1990) reported that crown diameter
and fruit yield increased 56% and 47%,
respectively, in an irrigated plot with
>70% soil moisture compared with a
nonirrigated plot with soil moisture of
50% to 60%. These reports suggest
that irrigation may be beneficial dur-
ing periods of extended drought, es-
pecially during flowering and fruit de-
velopment stages.
SOIL FERTILITY. Most of the soil
fertility research on sea buckthorn was
conducted in Russia and indicates that
sea buckthorn, like any other crop,
requires adequate soil nutrients for a
high yield of good-quality fruit. Sea
buckthorn responds well to phospho-
rus fertilizer, especially in soils low in
phosphorus. Application of 600 to 800
kg·ha–1 of calcium superphosphate
plowed deeply into the soil has been
recommended (A. Bruvelis, personal
communication). Garanovich (1995)
reported that, in Belarus, a single win-
ter top dressing with mineral fertilizers
100N–200P2O5–100K2O (kg·ha–1)
improved fruit size, yield, and quality.
Martemyanov and Khromova (1985)
indicated that best growth was ob-
tained by applying peat compost at 60
t·ha–1 and 50 kg·ha–1 each of N, P2O5,
and K2O. In Siberia, 5-year total fruit
yield increased by 23% when N, P2O5,
and K2O at 60 kg·ha–1 each were ap-
plied to a black calcareous soil
(Predeina, 1987). Montpetit and
Lalonde (1988) cautioned that nitro-
gen fertilization can adversely affect
root nodulation and it delays the de-
velopment of nodules after inocula-
tion with Frankia. Similar results have
been shown for other nitrogen-fixing
woody plants (Mackay et al., 1987).
Mishulina (1976) reported that foliar
sprays with micronutrients—Cu, Mo,
Mn, I, B, Co, and Zn—increased fruit
weight by up to 34.5%. In China,
application of 100 to 150 t·ha–1 of
compost or 400 to 500 t·ha–1 of green
manure is recommended before plant-
ing (Lu, 1992). Wolf and Wegert
(1993) noted that precise fertilizer
recommendations should be deter-
mined by soil sampling and analysis.
Fig. 2. The distribution of sea buckthorn
in Europe and Asia.

HortTechnology ● Oct./Dec. 1996 6(4) 373
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and Triquart, 1992) and Sweden (S.
Olander, personal communication).
WEED CONTROL. Weed control or
vegetation management is very impor-
tant in sea buckthorn plantings. Proper
weed control promotes growth of
newly planted seedlings. Cultivation
in new plantings should not disturb
the soil 8 to 10 cm below the surface so
that shallow roots are not damaged
(Gonchar and Saban, 1986). Herbi-
cides for sea buckthorn plantations
currently are being evaluated by the
PFRA Shelterbelt Centre. Several
chemicals are registered in Canada for
weed control in sea buckthorn
shelterbelts (Schroeder and Alspach,
1995). Albrecht et al. (1984) cautioned
that only low concentrations of herbi-
cides should be used. Low rates of si-
mazine and lenacil applied to cuttings of
sea buckthorn 10 days after planting
and again 2 months later gave effective
weed control and increased survival rate
and growth (Shlyapnikova, 1985).
MALE : FEMALE PLANT RATIO IN
THE PLANTING. For economical rea-
sons, the ratio of male to female plants
is important, as the number of fruit-
bearing trees should be maximized. In
Canada, sea buckthorn plants for
shelterbelts are grown from seeds. Con-
sequently, seedlings of unknown sex
are planted, which results in an uneven
distribution of male and female plants
within each planting. This practice has
long-term effects on total yield. To
avoid this problem, the easiest ap-
proach would be vegetative propaga-
tion from promising mature plants of
known sex. Recommendations for male
: female ratio vary. Gakov (1980) con-
sidered that 6% to 7% male trees is
adequate for pollination, whereas
Albrecht et al. (1984) and Wolf and
Wegert (1993) recommended 8% to
12%. The Siberian Institute of Horti-
culture in Russia recommended one
male : female mixed row for every two
rows of female plants. In the mixed
row, every fifth plant is a male.
Goncharov (1995) reported that this
design gave significantly higher fruit
yields compared to other designs. For
effective pollination, the male variety
should be cold resistant and have a
long flowering period, an adequate
amount of pollen, long-lived, and good
vigor (Garanovich, 1995), since a pol-
linator had an appreciable effect on
fruit weight, flavor, and ripening
(Buglova, 1981).
HARVESTING. Berries persist on
the branches all winter, perhaps due to
the absence of an abscission layer. This
results in an attractive ornamental plant
in winter, but it is undesirable for
harvesting. In Saskatchewan, Canada,
the total labor cost for harvesting an
orchard of 4 ha was estimated to be
58% of the total cumulative produc-
tion cost over 10 years. In Asia, har-
vesting is still done by hand or with
picking tools. This difficult and labor-
intensive process requires about 1500
h·ha–1 (Gaetke and Triquart, 1993).
Wolf and Wegert (1993) and Olander
(1995) reported that difficulties asso-
ciated with harvesting are major barri-
ers to orchard production of sea buck-
thorn in Europe. Therefore, one of the
most important factors for the success
of sea buckthorn as a viable cash crop
is a better and more economical har-
vesting method.
Koch (1981) harvested entire
fruiting shoots with pneumatic shears.
Botenkov and Kuchukov (1984) de-
veloped a device for hand-picking that
consists of two hinged jaws with teeth
and brushes. The development of
mechanical harvesting techniques for
sea buckthorn has attracted consider-
able attention. Savkin and
Mukhamadiev (1983) designed a prun-
ing machine to trim sea buckthorn
into a hedge suitable for mechanical
harvesting. Mechanical harvesters have
been developed in Sweden, Germany,
and Russia (Olander, 1995). How-
ever, most have disadvantages, such as
fruit and bark damage and low effi-
ciency. Harvesting equipment tested
include shakers (Affeldt et al., 1988;
Gaetke et al., 1991), vacuum suction
(Varlamov and Gabuniya, 1990), and
quick freezing (Gaetke and Triquart,
1993). Many harvesters are based on
the principle of cutting off fruit-bear-
ing branches (Gaetke and Triquart,
1992; Olander, 1995). Olander (1995)
reported development of an over-the-
row mechanical harvester that removes
fruit-laden branches and extracts the
fruit by shaking the branches in an
axial direction.
In Germany, considerable work
on mechanical harvesting of sea buck-
thorn has been completed. Wolf and
Wegert (1993) reported techniques in
which fruiting branches were removed
and frozen overnight at –36 °C. Fro-
zen fruit were removed by beating the
branches. This method provided ber-
ries with excellent quality, but it was
labor intensive (about 450 h·ha–1).
TEMPERATURE. Sea buckthorn can
endure the extreme minimum air tem-
perature of –43 °C without sustaining
long-term damage (Lu, 1992). Con-
versely, it can survive summer tem-
peratures up to 40 °C, although Lu
(1992) reported that temperatures
above 30 °C burned leaves on newly
planted seedlings. The latter observa-
tion may be the reason why efforts to
introduce sea buckthorn from moun-
tain areas to the plains of Asia often
have failed.
SPACING. The largest sea buck-
thorn population in North America is
in the Canadian prairies, where about
1000 km of field shelterbelts are planted
annually (Schroeder and Yao, 1995).
Two-year-old seedlings are planted for
shelterbelts normally in one to three
rows, 1 to 2 m apart within rows and 5
m between rows. Wolf and Wegert
(1993) recommended a spacing of 1 m
within the row and 4 to 4.5 m between
rows to allow equipment access, with
rows oriented in a north–south direc-
tion to provide maximum light.
Beldean and Leahu (1985) reported
that fruit yields are greatly influenced
by exposure to sunlight, as sea buck-
thorn will not tolerate shade. High
density orchards (1 × 1 m) are being
considered in Europe to facilitate over-
the-row harvesting equipment
(Olander, 1995).
PRUNING. The purpose of prun-
ing sea buckthorn is to train branches,
promote growth, and facilitate har-
vesting (Albrecht et al., 1984). Savkin
and Mukhamadiev (1983) reported
that moderate pruning will increase
the yield and fruiting life of the plants.
Sea buckthorn grows up to 2 to 3 m in
4 years and forms its crown at the base
of the main trunk. The crown should
be pruned annually to remove overlap-
ping branches, and long branches
should be headed to encourage devel-
opment of lateral shoots. In about the
fifth year, the main trunk stops grow-
ing, and branches begin to grow from
lateral buds. Mature, fruiting plants
should be pruned to allow more light
penetration if the bush is dense. To
prevent sea buckthorn from prema-
ture senescence, 3-year-old branches
should be pruned for rejuvenation (Lu,
1992). In Russia, pruning trials were
carried out with sea buckthorn with
the goal of creating a hedge to allow
efficient mechanical harvesting (Savkin
and Mukhamadiev, 1983). Similar
work is underway in Germany (Gaetke

HortTechnology ● Oct./Dec. 1996 6(4)374
The use of hormone treatments
to facilitate fruit release is promising.
Trushechkin et al., (1973) reported that
Ethrel (ethephon) at 2000 mg·L–1 of
water decreased fruit detachment force
by 30%. Demenko et al. (1986a) sug-
gested that the inability of ripe fruit to
Gaetke and Triquart (1993) devel-
oped a harvester that works on the
riddle principle. In this system, pruned
branches are hand fed into the har-
vester, and the fruit is separated from
branches and leaves by screen convey-
ors and fans.
Fig 3. Possible uses for components in
different sea buckthorn plant parts.

HortTechnology ● Oct./Dec. 1996 6(4) 375
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abscise is caused by compartmentation
of internal ethylene in seeds. Ethylene
treatment induced the formation of an
abscission layer, which could make
harvesting more efficient (Demenko
et al., 1986b; Demenko and
Korzinnikov, 1990).
YIELD. Like any other crop, yield
of sea buckthorn is affected by various
factors such as genotype, soil condi-
tions, annual precipitation, tempera-
ture, crop management, numbers of
fruit-bearing branches, and time and
methods of harvesting (Kondrashov,
1981). Yield data of sea buckthorn are
scarce, since most fruit collection is
from natural habitats, plantations for
controlling soil and water erosion,
and field shelterbelts. Wolf and Wegert
(1993) reported yield of 5 t·ha–1 from
an orchard plantation in Germany.
Total yields reported in the literature
from various regions were mostly esti-
mates from experimental plots, culti-
vated under ideal conditions and man-
agement (I.M. Garanovich, personal
communication). In Saskatchewan,
Canada, fruit production in
shelterbelts ranges from good to ex-
cellent, some selected plants produc-
ing 5 to 7 kg annually. This equates to
yields of 4 to 5 t·ha–1. Canada Sea
Buckthorn Ltd. (British Columbia)
estimated an average of 3.25 kg/tree
(6-year-old) collected from shelterbelt
plantings (C. McLoughlin,, personal
communication). Orchards with 4000
trees/ha and a 1:6 male and female
ratio should yield about 11 t·ha–1.
Detailed study is underway in Canada
on the effects of crop management on
yield.
Breeding and selection
There is a wide range of morpho-
logical diversity among sea buckthorn
seedlings and mature plants within
each subspecies, which is a good indi-
cation that there are excellent oppor-
tunities for plant improvement by
breeding or selection for desired char-
acteristics. Phenological observations
have shown a clear gradient of growth
rhythm and plant size corresponding
to geographical distribution (Yao and
Tigerstedt, 1994), which indicates that
selection based on responses of plants
to a specific growing region is impor-
tant. This selection procedure is im-
portant in any country before sea buck-
thorn can be developed into a new
viable crop.
Breeding sea buckthorn has been
conducted for decades in Russia
(Goncharov, 1995), Ukraine (Gladon
et al., 1994), China (Huang, 1995),
and Finland (Hirrsalmi, 1993). The
first breeding programs began with
mass selection from natural popula-
tions. This method is still common
practice (Wahlberg and Jeppsson,
1990) but is gradually being replaced
by hybridization (Huang, 1995; Yao
and Tigerstedt, 1994). Polyploid
breeding was reported in Russia where
autotetraploids were induced by colchi-
cine (Shchapov and Kreimer, 1988).
There have been no reports of genetic
engineering in sea buckthorn breed-
ing. Breeding programs exist in Swe-
den (Wahlberg and Jeppsson, 1990,
1992), Finland (Hirrsalmi, 1993; Yao
and Tigerstedt, 1994), and Germany
(Albrecht, 1993; Muller, 1993). Finn-
ish and Swedish programs have con-
centrated on mass selection and hy-
bridization. In Germany, the objective
was to establish a collection of sea
buckthorn representing the genetic
spectrum of the species (Muller, 1993).
A sea buckthorn breeding program
has been initiated recently in Canada
(Schroeder, 1995). It includes a long-
term breeding population made up of
progeny from diverse foreign wild col-
lections and a short-term breeding
population of superior individual plants
selected from local plantations.
Important characters that need
improvement in sea buckthorn are yield
(Kondrashov, 1986a; Huang, 1995),
fruit size (Buglova, 1978), winter har-
diness (Kalinina, 1987), thornlessness
(Hirrsalmi, 1993; Albrecht, 1993),
fruit quality and early maturity (Yao
and Tigerstedt, 1994), growth habit
and long pedicel for mechanical har-
vest (Wahlberg and Jeppsson, 1990),
and nitrogen-fixing ability (Huang,
1995). Kondrashov (1986a, 1986b)
reported that average fruit weight and
number of flower buds per unit length
of branch are the most reliable traits
for yield selection in the Altai region of
Siberia. P.L. Goncharov of Russia (per-
sonal communication) pointed out that
thornlessness could be selected in
young seedlings; however, yield of each
breeding line cannot be determined
until at least the fourth year. Fruit size,
thorniness, and hardiness reportedly
are controlled by quantitative genes,
and selection for one of these charac-
ters will not adversely affect others
(Huang, 1995).
Propagation
The most common methods for
propagating sea buckthorn are by seed,
softwood or hardwood cuttings, and
layering and suckers. Micropropaga-
tion using meristem culture has been
investigated (Burdasov and Sviridenko,
1988; Montpetit and Lalonde, 1988)
but is not commonly used.
SEEDS. Propagation from seed is
relatively simple and produces a large
number of seedlings at fairly low cost
compared with other propagation
methods. Storage affects seed viability.
Smirnova and Tikhomirova (1980)
reported that seeds of Hippophae
rhamnoides lost 60% of viability after 4
to 5 years of storage.
Internal seed dormancy can be
broken by stratification in moist sand
for 90 d at 5 °C (Siabaugh, 1974).
Seeds of sea buckthorn need high tem-
perature to germinate. At 10 to 12 °C,
Lu (1992) reported 13.2% germina-
tion after 47 d compared to 95% in 6 d
for seeds at 24 to 26 °C. Vernik and
Zhapakova (1986) also reported more
rapid germination at 25 to 27 °C than
at 20 °C. Good results also can be
obtained by soaking seeds in hot water
(70 °C) for 24 to 48 h, stirring fre-
quently, then letting the water cool to
room temperature (Lu, 1992). After
soaking, seeds should be air dried be-
fore sowing. This technique is useful
when sowing outdoors in spring or
indoors in a greenhouse. At the PFRA
Shelterbelt Centre, Saskatchewan,
nonstratified seeds sown in late Sep-
tember at a depth of 1 cm and a rate of
100 seeds/m in rows 60 cm apart
emerged the following spring with a
90% germination rate.
CUTTINGS. Cuttings produce
rooted plants with the same genotype
as the parent plant. The cuttings will
bear fruit 1 to 2 years earlier than seed-
propagated trees. Sea buckthorn can
be propagated using either hardwood
or softwood cuttings.
HARDWOOD CUTTINGS. The per-
centage of successful rooting from
hardwood cuttings varies. Avdeev
(1984) reported 86% to 100% success.
Garanovich (1984) reported that, in
the greenhouse under artificial mist,
rooting success was 20% lower with
hardwood cuttings than with softwood
cuttings, but plants from hardwood
cuttings attained heights of 90 cm by
the end of the first growing season and
could be planted out the next spring,

HortTechnology ● Oct./Dec. 1996 6(4)376
whereas plants from softwood cut-
tings needed 1 to 2 years before trans-
planting to the field. Kondrashov and
Kuimov (1987) reported that hard-
wood cuttings with apices removed
were rooted successfully outdoors un-
der plastic in pure sand or a sand and
peat (1:1) mixture. In a separate ex-
periment, they showed that 2-year-old
wood, cut before budbreak and stored
for 10 d in sawdust at 10 to 15 °C, gave
100% rooting in the field. Kuznetsov
(1985) recommended taking cuttings
before bud break, soaking in water (18
to 20 °C) for 7 days, and planting in
the field with dark polyethylene mulch.
In British Columbia, we obtained 90%
rooting of cuttings taken in mid-March,
stored in plastic bags at 0 °C until May,
and placed in pots filled with peat in a
heated propagation box (18 to 22 °C)
indoors under fluorescent light. Lu
(1992) reported that hardwood cut-
tings have not been used widely in
nurseries.
SOFTWOOD CUTTINGS. Avdeev
(1976) reported that softwood cut-
tings collected from an 8-year-old tree
in early spring, treated with IBA (50
mg·L–1) solution and planted in a peat
and sand (2:1) mixture under mist,
had 96% to 100% rooting in 9 to 11 d.
Balabushka (1990) compared the ef-
fects of IAA, IBA, and chloro-
phenoxyacetic acid on rooting of sea
buckthorn and found that IAA root
dips were superior to the other hor-
mone treatments. Ivanicka (1988) re-
ported that, with or without IBA (0.1%
to 0.3%) treatment, semi-lignified
Hippophae rhamnoides cuttings rooted
readily in a peat, polystyrene granules
and sand (1–2:1 :0.5) mixture under
mist in a plastic house. Timing of
cutting collection is important. Kniga
(1989) reported that, in the Kiev re-
gion of Ukraine, the optimum time
was late May. Kondrashov and Kuimov
(1987) reported that cuttings taken in
late June from severely pruned branches
(pruning was conducted in early spring
before budbreak) successfully rooted
(95% to 98%) in the greenhouse under
mist.LAYERING AND SUCKERS. Root
cuttings also can be an effective propa-
gation method for sea buckthorn (A.
Bruvlis of Latvia, personal communi-
cation). Root cuttings were planted in
pots and placed in a greenhouse for 6
weeks before being transplanted to the
field at a spacing of 8 × 20 cm. Cut-
tings need to be acclimated to field
conditions before planting by placing
pots in a shady area for 1 week. The
best results were obtained in sandy
loam at pH 6 to 6.5 with medium
humus content. Sea buckthorn easily
produces suckers within a few years of
planting, which is a good source for
propagation (Kondrashov and Kuimov,
1987). The possibility of invasiveness
by suckers to surrounding areas is high;
routing cultivation and herbicide ap-
plication are the best control measures
for this potential weediness character-
istics of sea buckthorn.
Multipurpose use
ENVIRONMENTAL VALUE. The wide
adaptation, fast growth, strong
coppicing, and suckering habits,
coupled with efficient nitrogen fixa-
tion, make sea buckthorn particularly
suitable for planting in degraded soils.
In China, sea buckthorn has controlled
soil erosion and water loss effectively
and increased land reclamation. In
addition, the harvest of sea buckthorn
fruit has provided value-added indus-
tries to support the economy of rural
regions of Asian countries. In Canada,
sea buckthorn has proved highly ben-
eficial for enhancement of wildlife habi-
tat, farmstead shelterbelts, erosion con-
trol, and mineland reclamation.
CHEMICAL COMPONENTS. Sea
buckthorn fruit is rich in carbohy-
drates, protein, organic acids, amino
acids, and vitamins (Bernath and
Foldesi, 1992). Fruit contain 16 to 28
mg carotenoids/100 g fruit, which
can be used as food additives
(Kudritskaya et al., 1989). Flavonoid
content in leaves and fruit ranges from
310 to 2100 mg/100 g air-dried leaf
and 120 to 1000 mg/100 g fresh fruit,
respectively (Chen et al., 1991;
Glazunova et al., 1984, 1985). Total
volatile oil from the fruit is 36 mg·kg–1
(Hirvi and Honkanen, 1984), dry
matter is 24.6% to 33.8% (Igoshina et
al., 1987), and essential oil extracted
from seeds ranged from 8% to 12%
(w/w) (Lu, 1992).
Bounous and Zanini (1988)
found that fruit maturity affects N, Ca,
K, Na, Mg, Cu, Fe, Zn, Mn, titratable
acidity, pH, moisture, glucose, fruc-
tose, and ascorbic acid content. Harju
and Ronkainen (1984) reported that
the trace elements found in liqueurs
prepared from sea buckthorn included
Al, As, Ca, Cd, Cr, Cu, Fe, K, Mg, Mn,
Na, Ni, Pb, Rb, and Zn. Differences in
chemical composition, carbohydrates,
moisture, lipids, acids ,and vitamins A
and C were found among the large and
small fruit (Chen et al., 1991) and
among subspecies (Lu, 1993) and geo-
graphic locations (Wang, 1990).
Eliseev (1976) indicated that the levels
of the biologically active substances
such as ascorbic acid and carotene
were higher in tree-like forms than in
bush forms. Franke and Muller (1983)
analyzed the fat of fruit and seeds and
found 47%, 21% saturated fatty acids
and 53%, 39% unsaturated fatty acids
respectively.
NUTRITIONAL VALUES. The value
of sea buckthorn is often based on the
nutritional value of its fruit (Magherini,
1986). Sea buckthorn berries are
among the most nutritious and vita-
min-rich fruit known. The fruit, in-
cluding seeds, contain a large amount
of essential oils and vitamin C
(Centenaro et al., 1977; Novruzov
and Aslanov, 1983). The vitamin C
concentration in berries varies depend-
ing on species, geographical location,
and physiological maturity (Bernath
and foldesi, 1992; Zhou et al., 1991)
from 360 mg/100 g of berries for the
European subspecies rhamnoides
(Rousi and Aulin, 1977; Yao et al.,
1992) to 2500 mg/100 g of berries
for the Chinese subspecies sinensis (Yao
and Tiherstedt, 1994), which is higher
than strawberries (205 mg vs. 64 mg/
100 g; Gontea and Barduta, 1974),
kiwi fruit (300 to 1800 mg vs. 100 to
470 mg/100 g), orange (50 mg/100
g), and tomato (12 mg/100 g) (Lu,
1992). Sea buckthorn is also high in
protein, especially globulins and albu-
mins (Solonenko and Shishkina, 1983),
carotene (Kostyrko, 1990), fatty acids
(Loskutovaet al., 1989), and vitamin
E (Bernath and Foldesi, 1992). Fatty
acids and vitamin E content are higher
than in wheat, safflower, maize, or
soybean (Lu, 1992). The leaves of sea
buckthorn contain many nutrients and
bioactive substances, such as urtica
dioica, vaccinium myrtilis, and berberis
vulgaris, which are suitable for animal
feed (Morar et al., 1990).
MEDICINAL VALUE. Medicinal uses
of sea buckthorn are well documented
in Asia and Europe. Clinical investiga-
tions on medicinal uses were initiated
in Russia during the 1950s (Gurevich,
1956). Sea buckthorn oil is approved
for clinical use in hospitals in Russia
and in China, where it was formally
listed in the Pharmacopoeia in 1977
(Xu, 1994). More than 10 different

HortTechnology ● Oct./Dec. 1996 6(4) 377
RR
RR
REVIEWEVIEW
EVIEWEVIEW
EVIEW
drugs have been developed from sea
buckthorn in these countries and are
available in different forms (e.g., liq-
uids, powders, plasters, films, pastes,
pills, liniments, suppositories, aerosols,
etc.) and can be used for treating oral
mucositis, rectum mucositis, vaginal
mucositis, cervical erosion, radiation
damage, burns, scalds, duodenal ul-
cers, gastric ulcers, chilblains, skin ul-
cers caused by malnutrition ,and other
skin damage (Abartene and
Malakhovskis, 1975; Buhatel et al.,
1991; Chen, 1991; Cheng et al., 1990;
Dai et al., 1987; Kukenov et al., 1982).
The most important pharmacological
functions of sea buckthorn oil can be
summarized as diminishing inflamma-
tion, disinfecting bacteria, relieving
pain, and promoting regeneration of
tissues. It also can be used for skin
grafting, cosmetology, and treatment
of corneal wounds. In an unconfirmed
report from China, 350 patients treated
with beauty cream made from sea buck-
thorn oil had positive therapeutic ef-
fects on xanthopsis, melanosis, senile
skin wrinkles, and freckles (Zhong et
al., 1989). Russian research reported
that 5-hydroxytryptamine (hippophan)
isolated from sea buckthorn bark in-
hibited tumor growth (Sokoloff et al.,
1961). More recently, studies on the
anti-tumor effects of sea buckthorn oil
have shown positive results in China
(Zhong et al., 1989).
Conclusion
Sea buckthorn is a unique and
valuable plant species currently being
domesticated in various parts of the
world. The species has been used to a
limited extent in North America for
conservation plantings, but the use of
food and nonfood sea buckthorn prod-
ucts has not been pursued. The plants
are easily propagated and yields are
relatively high, and production is reli-
able, with the potential market mainly
in Europe. The main constraint to
large-scale fruit production in North
America is harvesting. This problem is
being addressed through breeding pro-
grams and equipment development.
Most sea buckthorn research has been
conducted in Asia and Europe, specifi-
cally China, Russia, and Germany.
Recently interest in western Europe
and Canada has increased, and active
research programs are underway.
Unique plant products, especially
those with proven nutritional quality,
are gaining popularity in North
America. Development of a North
American sea buckthorn industry pre-
sents a unique opportunity for agricul-
tural production of a value-added crop
on marginal land.
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