Scientia Horticulturae 205 (2016) 7078

Contents lists available at ScienceDirect

Scientia Horticulturae
journal homepage: www.elsevier.com/locate/scihorti

Review

Potentials in Prunus mahaleb L. for cherry rootstock breeding

K. Hrotk
Department of Floriculture and Dendrology, Faculty of Horticultural Science, Szent Istvn University, P.O. Box: 1518 Budapest Pf. 53, Hungary

a r t i c l e

i n f o

Article history:
Received 1 February 2016
Received in revised form 1 April 2016
Accepted 12 April 2016
Available online 26 April 2016
Keywords:
Adaptability
Alkaline soil
Clonal rootstocks
Drought
Graft compatibility
Precocity
Propagation
Rootstock vigor
Seed trees

a b s t r a c t
Overview on Prunus mahaleb L. rootstock breeding and evaluation compile the knowledge on variability,
rootstock traits based on systematic evaluation and results achieved in previous breeding projects. Conclusion is that adaptability of mahaleb cherry to continental climate, tolerance to drought, hot summer,
poor soils, high pH and lime tolerance are valuable rootstock traits considering the forecasted environmental conditions due to the climate changes. The diverse rootstock usage in vigor control is the present
situation in the sweet and sour cherry growing. This diverse rootstock usage might be continued for
the future too, when training system and rootstock are considered together, matched properly with the
vigor of orchard site. There are few clonal rootstocks showing vigor 6570% compared to seedlings with
medium precocity. Further on the potential in compact or spur type mahalebs, genetic dwarf genotypes or
inbred lines are still not fully utilized in mahaleb cherry breeding. Possible interspecic crosses between
P. mahaleb and other cherry species (Prunus avium and Prunus fruticosa crosses are known yet) also could
extend the range of desired growth control and precocity. Traditional seed tree selection resulted in
seed tree clone, producing high quality seedlings for sweet- and sour cherry growing but their vigor
control and precocity does not meet all the requirements of modern pedestrian orchards. Variability
of mahaleb cherry manifested in selected genotypes, inbred lines and potential of interspecic crosses
promise progress in those directions that are desired rootstock traits in vigor control, precocity and
environmental adaptability.
2016 Elsevier B.V. All rights reserved.

Contents
1.
2.
3.
4.

5.

6.

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Botany and classication . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Utilization of P. mahaleb as cherry rootstock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Important rootstock traits of P. mahaleb L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
4.1.
Graft compatibility . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
4.2.
Rootstock vigor, precocity and productivity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
4.3.
Propagation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
4.4.
Training system and pruning . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
4.5.
Environmental adaptability . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
Achievements of P. mahaleb breeding as cherry rootstock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
5.1.
Seed tree selection and establishing of seed orchards . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
5.2.
Inbreeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
5.3.
Clonal rootstock selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
5.4.
Interspecic hybrids of P. mahaleb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76

E-mail addresses: hrotko.karoly@kertk.szie.hu, karoly.hrotko@gmail.com

http://dx.doi.org/10.1016/j.scienta.2016.04.015
0304-4238/ 2016 Elsevier B.V. All rights reserved.

K. Hrotk / Scientia Horticulturae 205 (2016) 7078

1. Introduction
Major cherry growing countries used in large ratio Prunus
mahaleb (L.) as cherry rootstock, but very little breeding work has
been done to improve its rootstock traits. Adaptability of mahaleb
cherry to continental climate, tolerance to drought, hot summer,
poor soils and lime will increase its importance in the future cherry
growing. These rootstock traits may contribute to improve adaptability of cherry growing to those environmental conditions that
due to the climate changes are usually forecasted. Since the expectations of growers and researchers in the rst half of the last century
considering the growth control by usage of mahaleb cherry were
not fullled, breeders showed few attention to P. mahaleb L.
Albeit the modern orchard systems, the so called pedestrian
orchards of sweet cherry requires dwarng and precocious rootstocks, the rootstock usage in sweet and sour cherry orchards is still
diverse. No doubt that training system and rootstock must be considered together, matched properly with the vigor of cultivar and
orchard site involving soil fertility and climate. Growers for intensive orchards producing hand-picked cherries for fresh market
show increased interest for dwarng rootstocks, which allow plantation density up to 10005000 trees/ha (Robinson, 2005; Sansavini
and Lugli, 2014; Musacchi et al., 2015). On the other hand several training systems with special training and pruning protocols
(KGB, Spanish bush, Steep leader, Tall spindle axe, UFO) allow establishing pedestrian or semi-pedestrian orchards on semi vigorous
and vigorous rootstocks (Robinson, 2005; Negueroles, 2005; Ercisli
et al., 2006; Iglesias and Peris, 2007; Hrotk, 2010; Lang, 2011; Long
et al., 2015). The growth controlling effect of dwarng rootstocks in
these case could be partly replaced by application of special training and pruning protocol, frequent summer pruning, root pruning
and water restriction (Negueroles, 2005; Iglesias and Peris, 2007;
Hrotk, 2010; Lang, 2011; Long et al., 2015).
Recently some researchers reports show increasing interest on
P. mahaleb breeding or selection of promising genotypes. The accumulated experience on mahaleb cherry breeding and evaluation as
cherry rootstock could be worthy pool of information for breeding
in mahaleb cherry, and show the potentials and possible ways for
the future breeding programs.

2. Botany and classication

By systematic classication of cherry species according to
Rehder (1990) P. mahaleb L. belongs to the III. Cerasus subgenus,
Section 5 Mahaleb Focke, which section among others involves the
species P. mahaleb L. native to Europe and Western-Asia. The geographic distribution of mahaleb cherry spread from Central- and
South-Europe, Northern-Africa through S-E-Europe, Asia Minor,
Western and Central Asia (Turkey, Iraq, Iran, Kyrgyzstan; Tajikistan;
Turkmenistan) and probably some provinces of NW-China.
P. mahaleb occurs in thickets and open woodland on dry slopes;
in central and southern Europe typical in thickets on dry karst
areas (Terp, 1968; Faust and Surnyi, 1997). The origin of name
mahleb or mahlab is Arabian, in Arabian speaking countries
domesticated in ancient ages and grew as medicinal plant and spicy.
It is still used as food additive, medicinal plant and spicy in Asia
Minor and Western-Asia (Ozturk et al., 2014). In mahaleb cherry
fruit valuable phenolics, phenolic acid derivatives, quercetin glycosides, anthocyanins and coumarin were characterized (Ieri et al.,
2012). Bark, wood, leaves and seeds contain coumarin providing a
fragrant smell to the plant and its seeds. Pipe manufacturer prefer
mahaleb wood for pipe and pipe stem. Useful revegetator plant, few
cultivated varieties have been selected for their ornamental value,
including the compact Monstrosa (var. bommii, var compacta)
Pendula (var. pendula), with drooping branching, and Xantho-

71

carpa (var. chrysocarpa) with yellow fruit (Krssmann, 1978a,b).

In most of the cherry growing countries mahaleb cherry is used as
rootstock for both sweet and sour cherries.
The variability of P. mahaleb (Krssmann, 1978b; Hrotk, 1996;
Ganji and Khalighi, 2006; Shahi-Gharahlar et al., 2011; KhadiviKhub et al., 2012; Koc and Bilgener, 2013) is still not fully utilized
in rootstock breeding. P. mahaleb occurs in great diversity and has
been classied by Terp (1968) into several sub-species native
to different geographic areas and environmental conditions. From
among them ssp. mahaleb is known as small-leaved mahaleb, distributed in Western Europe in more mild and humid climates,
characterized by small leaves and ne pubescent young shoots.
This is hardy in Europe, but the long growing period to late autumn
makes it sensitive to early frost injuries under continental climate.
The geographic area of ssp. simonkaii (P<nzes) Terp (known as
broad-leaved mahaleb cherry, leaves are large, shoots are hairless)
distributes from Central-Europe through Asia-Minor, Western and
Central Asia to China. This subspecies is more adapted to continental climate, shoot growth terminates earlier, so it is completely
winter-hardy in temperate zone conditions. The areal of third subspecies, ssp. cupaniana (Guss) Terp is located in south-eastern
Europe and Asia Minor, leaves are leather-like, young shoots are
hairless and fruits are little larger.
3. Utilization of P. mahaleb as cherry rootstock
Usage of mahaleb cherry as rootstock vary by countries depending on the site conditions and the rate of sour cherries among the
total cherry production. Sour cherry growers in majority of countries prefer P. mahaleb (L.) as rootstock but it is applied for sweet
cherry too in considerable rate in such large sweet cherry growing
countries like Turkey (Ercisli et al., 2006), Iran (Ganji and Khalighi,
2006), China (Faust et al., 1998; Cai et al. 2007; Hrotk and Cai,
2014) and United States (Webster and Schmidt, 1996; Lang 2000,
2006; Kappel et al., 2005; Long et al., 2005, 2015). From among the
Central-European countries in Poland (Rejman et al., 2002; Barya
and Kapan, 2005; Stachowiak et al., 2014), in Eastern provinces
(countries) of Germany (Balmer and Blanke, 2005; Balmer, 2008)
and in Hungary (Hrotk, 2008) is planted in considerable rate as
cherry rootstock. In South- and South-East Europe P. mahaleb is
considered as traditional rootstock in Spain (Moreno et al., 1996;
Negueroles, 2005; Iglesias and Peris, 2007), in Southern-France
(Claverie, 1996; Edin et al., 1996; Saunier and Claverie, 2001), in
Italy (De Salvador et al., 2005), in Serbia (Milosevic et al., 2014)
in Bulgaria (Koleva, 2001; Sotirov, 2005; Lichev and Papachatzis,
2008, 2009) in Macedonia (Kiprijanovski and Gjamovsky, 2015)
and in Greece (Lichev and Papachatzis, 2009) and mahaleb cherry
as rootstock is still planted in considerable or major rate of cherry
orchards. In East-European countries (Ukraine and Russia) the large
rate of sour cherry orchards makes mahaleb cherry for major rootstock. In countries, where the P. mahaleb (L.) is used as rootstock
for sweet and sour cherries, the rootstock selection produced in the
last century seed-tree genotypes and seed orchards. The seedlings
in standard vigor are valuable rootstocks in continental climate and
certain soil conditions. The vegetative propagated clonal P. mahaleb
rootstocks, except for Sainte Lucie 64 and, are still in evaluation or
introduction stage.
4. Important rootstock traits of P. mahaleb L.
Rootstock breeding is a particularly long term endeavour and
progress tends to be slow regardless of which method used to
develop the candidate rootstocks. At the moment pre-selection
techniques are not available. Therefore to determine if the selection has any potential the candidate rootstock will need to be

72

K. Hrotk / Scientia Horticulturae 205 (2016) 7078

Table 1
Selected seed tree clones of Mahaleb cherry.
Variety name

Origin and attributes

Reference

Heimann X.
Alpruma

Germany, I1 progeny of self-fertile tree

Germany, Altenweddingen, seed mixt. of clones AF 5/19, AF 3/9, AF
6/16 and PB 9, pollinating each other, standard vigor
Hungary, Cegld, seed tree clone C500, pollinated by C2753 and rdi
V., standard vigor
Hungary, Cegld, seed tree clone C2753, pollinated by C500 and rdi
V., standard vigor
Hungary, Cegld, seed tree clone rdi V., pollinated by C500 and
C2753., standard vigor
Hungary, Budapest; I1 progeny of Korponay self-fertile seed tree, vigor
80% of standard
France, Bordeaux, I1 progeny of INRASL 405 self-fertile seed tree
USA, MI; Seedling of Nos Mahaleb 902, 904, 908, 916, standard vigor
Moldavia, selection from natural forest stands, produce standard vigor
seedlings. Pollination and mating system is not known

Heimann (1932)
Funk (1969)

Ukraine, selection from natural forest stands, produce standard vigor

seedlings. Pollination and mating system is not known
Bulgaria, selection from natural forest stands, produce standard vigor
seedlings. Pollination and mating system is not known
Bulgaria, selection from natural forest stands, produce semi vigorous
seedlings. Pollination and mating system is not known
Poland, selection from natural forest stands, produce semi vigorous
seedlings. Pollination and mating system is not known
Poland, selection from natural forest stands, produce semi vigorous
seedlings. Pollination and mating system is not known

Tatarinov and Zuev (1984)

CEMA
CEMANY
rdi V.
Korponay
FerciPontaleb
Mahaleb 900
Rozovaya prodolgovataya,
Chernaya Kruglaya iz
Bykovtsa, Nr 1 iz
Solonchen
Mahaleb N 24
IK-M 8
IK-M 9
Piast
Popiel

propagated to obtain sufcient plant material to provide for an

adequate eld test. Then a range of cultivars will need to be propagated on these candidates for eld evaluations. A number of years
are required to determine if there is any effect on scion precocity,
vigor, yield, and fruit quality. However at each stage (multiplication,
scion propagation etc.) unsuitable candidates can be discarded. For
example if propagation is extremely difcult or costly for a particular candidate, it can be discarded.

4.1. Graft compatibility

Graft compatibility is the major determining rootstock trait, on
which for mahaleb cherry contradicting opinions are known in
the literature. Graft incompatibility occurs when the composite
tree is produced from two or more species, so mahaleb cherry as
rootstock may involve some incompatibility with sweet and sour
cherries, based on several differences in their metabolic system.
This is well demonstrated by Favr-Bonvin et al., 1968 on example
of coumarin metabolism. Shoot system and roots of ungrafted P.
mahaleb tree transform phenyalanin into herniarin and coumarin.
Prunus avium is not able to synthesize these compounds or what
is more, may block the enzyme forming coumarin in mahaleb
shoot system when grafted onto P. avium root. Feucht (1982) supposed competitive inhibition in the background. In contrary, the P.
avium shoot system does not block the coumarin formation in P.
mahaleb root. So the metabolic system of heterospecic grafts is
more or less stressed (Feucht, 1982; Treutter et al., 1993; Feucht
et al., 2001), this is the case for cherry trees on P. mahaleb. Incompatibility symptoms may not manifest themselves under optimal
growth conditions; however when the tree is overcome by environmental stress the underlying incompatibility will be revealed.
Early data of Schnberg (1963), Garner (1967) and de Haas and
Hildebrandt (1967) suggest that the incompatibility of mahaleb
cherry occurs when the site is not properly selected for the rootstock and/or the scion. This could be the reason of the several
contradicting statements on the compatibility of mahaleb cherry.
Sour cherry cultivars usually show good compatibility on P.
mahaleb.

Nyjt (1987)
Nyjt (1987)
Nyjt (1987)
1968 and (Hrotk, 2004)
Sebok
Claverie (1996)
Perry (1987)
Yoltuchovski (1977) and
Tatarinov and Zuev (1984)

Koleva (2001) and Sotirov

(2005)
Koleva (2001) and Sotirov
(2005)
Rejman et al., 2002; Barya and
Kapan, 2005
Rejman et al., 2002; Barya and
Kapan, 2005

Mahaleb seedlings of selected seed-trees and clonal mahalebs

are tested and proved to be compatible with major sweet cherry
cultivars under optimal pedoclimatical conditions (Baryla and
Kaplan, 2005; Bujdos and Hrotk, 2014; Hrotk et al., 1999,
2009a,b; Sitarek et al., 1998; Sotirov, 2005, 2012, 2015; Stachowiak
et al., 2014; Stachowiak et al., 2015). Several examples demonstrate that properly selected mahaleb cherry as rootstock can give
compatible, long living and productive composite tree with both
sweet and sour cherries (Perry, 1987; Webster and Schmidt, 1996).
Incompatibility symptoms can include: poor bud take; the scion
snapping off at the bud union; small yellow leaves; stunted growth;
early reddening and fall of leaves in the autumn; scion or rootstock overgrowth; excessive rootstock suckering; and excessive
early fruiting (Perry, 1987; Webster and Schmidt, 1996).

4.2. Rootstock vigor, precocity and productivity

Very few information is available on the mechanism of vigor
control in cherry trees. One major components are the hormonal
interactions between rootstock and scion similarly to other composite trees (Feucht, 1982; Faust, 1989; Treutter et al., 1993;
Webster, 1998; Hrotk, 2008). However many elements of growth
control can be explained by hormonal control, the whole complex,
involving the rootstock effects on assimilate partitioning, water
and nutrient uptake and translocation, the mechanism of rootstock
effect on precocity and cropping efciency need further intense
research (Tanrisever and Feucht, 1978; Faust, 1989; Webster, 1998;
Hrotk, 2008). Misirli et al. (1996) related the vigor of the tree to
sieve tube size when selecting for low vigor Mahaleb rootstocks.
They found a direct relationship between vigor and the size of sieve
tube elements in wood of old trees, but no correlation in young trees
therefore they concluded that it cannot be used as a criterion for
predicting vigor. Goncalves et al. (2007) found signicantly higher
vessel frequency, lower vessel diameter, and relative hydraulic conductivity in dwarfed trees, especially grafted on Gisela 5 than trees
on the invigorating rootstocks, P. avium L., CAB 11E and Maxma 14.
Similarly higher vessel frequency and lower vessel diameter was

K. Hrotk / Scientia Horticulturae 205 (2016) 7078

73

Table 2
Vegetative propagated P. mahaleb rootstocks and derivatives (P. mahaleb as female parent).
Variety

Origin and attributes of clonal mahaleb rootstocks

References

SL 64

Selected in France from wild genotypes, vigorous, easy to propagate,

good compatibility and productivity with sweet and sour cherries.
Rooting of softwood cuttings in 95%

Bonn 60
Bonn 62
Bogdny

Vigorous clones selected in Germany, did not get into commercial

propagation
Hungary, selected as root sucker of an old and productive sweet cherry
tree, vigorous, wide crotch angles, good compatibility and
productivity. Rooting of softwood cuttings in 80%
Moderate vigorous selected in Hungary, good compatibility and
productivity, wide crotch angles. Rooting of softwood cuttings in 95%
Semi-vigorous clone, selected in Hungary,
good compatibility and productivity, wide crotch angles. Rooting of
softwood cuttings in 70%
Bulgaria, semi vigorous, rooting of softwood cuttings 50%

Thomas and Sarger (1965),

Claverie (1996), Edin et al.,
(1996) and Hrotk et al.
(1999)
Baumann, 1977

Egervr
Magyar

IK-M9
Mahaleb 2086
UCMH 55
UCMH 56
UCMH 59
Interspecic hybrids. P. mahaleb P. avium
MxM 2
and
MxM 60
MxM 14
and
MxM 97
OCR 2 and OCR 3

German mahaleb selection, Institute Dresden-Pillnitz, propagated by

softwood cuttings, semi-vigorous, tested in Bulgaria
USA, Calif. selection from open pollinated seedlings, standard vigor,
Phytophtora tolearant
USA, Calif. selection from open pollinated seedlings, standard vigor,
Phytophtora tolearant
USA, Calif. selection from open pollinated seedlings, semi-vigorous,
Phytophtora tolearant
USA, Oregon, very vigorous, adaptability like on Mahaleb, good
compatibility, resistant to Phytophtora, narrow crotch angle, more
precocious than seedling, good productivity
USA, Oregon, moderate vigorous, adaptability like on Mahaleb, good
compatibility, resistant to Phytophtora, narrow crotch angle, more
precocious than seedling, good productivity
USA, Oregon, vigorous

found in stem cross section of SL 64 compared to the semi vigorous

MxM 14 (Vgvri et al., 2008).
Vigor of mahaleb seedlings differ from Mazzard (P. avium)
seedlings: growth dynamics of trees on mahaleb cherry is faster
in early years, turning to bearing is earlier, than the growth slow
down by the 7th leaf, while on mazzard the growth of the tree in
later years is still strong (Perry 1987; Hrotk, 2004, 2008). The vigor
of seedling progeny of cross pollinated seed trees (most of the seed
trees are self-sterile) show typical hybrid vigor; the tree size on
such seedlings is similar to Mazzard. Trees on seedlings of certain
self-fertile seed trees (e.g. Korponay) showed 2025% vigor control (Hrotk 1990, 2004; Bujdos and Hrotk, 2014) compared to
SL 64. The seedling progeny even of semi-vigorous clones with selfsterile owers, coming from cross pollination show usually hybrid
2006). This is the case of mental stress
vigor (Hrotk and Erdos,
the underlying incompatibility will be revealed. Early dataMagyar
clonal rootstock, which is semi vigorous when propagated by softwood cuttings, while its seedlings are vigorous (Hrotk, 1990, 2004;
Bujdos and Hrotk, 2014). So it is necessary to evaluate distinctly
the seedling progeny and the clonally propagated rootstocks. Further inbreeding reduces the vigor of seedlings: I2 seedling progeny

of I1 inbred lines of Korponay trees as rootstock of rdi boterm

o
sour cherry cultivar showed 50% vigor control compared to the
seedlings of Korponay (I1 population) (Hrotk and Magyar, 1998;
Hrotk et al., 2005).
An extended vigor range from standard to medium is found
in clonal P. mahaleb (Hrotk, 2004; Hrotk and Magyar, 2004;
Lang, 2006; Sotirov, 2012; Stachowiak et al., 2014; Barac et al.,
2014); their vigor control is about 2035% compared to standard
vigor seedlings. As registered semi vigorous rootstocks followings
are available: IKM 9 (Koleva, 2001), Magyar (Hrotk, 2004) and
UCMH 59 (Lang, 2006). Further promising selections are reported
by Sotirov (2012), Stachowiak et al. (2014) and Barac et al. (2014).
Selection of P. mahaleb in Iran (Ganji and Khalighi 2006; Shahi-

Hrotk (2004), Hrotk et al.

(2009c) and Magyar and
Hrotk (2013)
Hrotk (2004) and Bujdos
and Hrotk (2014)
Magyar and Hrotk (2013)

Koleva (2001) and Sotirov

(2005)
Sotirov, 2012
Lang (2006)
Lang (2006)
Lang (2006)

Westwood (1978) and

Perry (1987)
Westwood (1978), (Perry
1987), Edin et al. (1996)
and Hrotk et al. (1999)
Perry (1987)

Gharahlar et al., 2011) and in Turkey (Koc and Bilgener, 2013)

resulted wide range of morphology and vigor of mahaleb trees, but
no further information is available on the intended seed or clonal
propagation and evaluation results as rootstocks. From Poland
Stachowiak et al. (2014, 2015) report on evaluation of Mahaleb
biotypes Nr. 2 and Nr. 6, which showed lower vigor and higher
productivity compared to Mazzard.
In the 19th and in the rst half of the 20th century growers and
researchers expected vigor control when using mahaleb as rootstock (Kppers, 1978) but these expectations were not fullled by
usage of mahaleb cherry. Testing of clonal and seedling mahaleb
rootstocks proved that no denite dwarng effect has been found
in P. mahaleb genotypes, although the compact or genetic dwarf
mahaleb genotypes are signs that there are some sources for vigor
control in this species (Krssmann, 1978b; Jacob, 1992; Hrotk and
Magyar, 1998; Hrotk, 2004; Hrotk et al., 2005). Further on the
possible utilization of inbred lines (Hrotk et al., 2005) has not
been fully investigated. Despite of the missing vigor control further breeding efforts are needed to increase the vigor range, and
improve the precocity of P. mahaleb (L) as cherry rootstock. Genetic
diversity could be increased by inbreeding, as well as crosses with
genetic dwarf genotypes and interspecic hybridization, which is
followed by clonal selection.
Branch angle of scion can also be affected by rootstock. Hrotk
et al. (1999, 2008) observed that scions on P. mahaleb Magyar
showed larger crotch angle, while on MxM 14 and MxM 97 the
crotch angle of sweet cherry trees was narrower.
Precocity, abundance and consistency of yield as well as fruit
quality are affected by rootstocks, but there is considerable interaction between rootstock, training and pruning, tree spacing and
nutrition. Perry (1987) reported that scions on Mahaleb seedling
rootstock produced fruit one to two years earlier than on Mazzard
rootstocks, this is conrmed by (Hrotk, 1990), and Stachowiak
et al. (2014, 2015) in several trials. Intensive orchards with close

74

K. Hrotk / Scientia Horticulturae 205 (2016) 7078

spacing of trees and fruiting wood management can also contribute

to precocity (Hrotk et al., 2009a,b,c).
Scion productivity can only be evaluated with eld trials. A relationship between yield efciency, crop load and leaf area can affect
fruit size. Concerning the fruit size and other quality attributes
some trials (Edin et al., 1996; Simon et al., 2004; Cittadini et al.,
2007; Gyeviki et al., 2012) showed relations between yield efciency, crop load and leaf area. Since highly efcient dwarng
rootstocks extremely increase the fruit number to leaf area, this
leads to diminishing of fruit size and quality, while on medium
vigorous mahaleb rootstock the cherry cultivars still produce large
fruit size, keeping the optimal fruit to leaf area ratio.

4.3. Propagation
Propagation by seed is most simple way for producing rootstock liners. For the successful seedling production the breaking
dormancy, uniform germination, high germination rate and germination speed is essential. In nursery practice both fresh harvested
and one-year-old stored seed is used, but the improving the germination rate and velocity is essential at both seed age (Krssmann,
1978a; Hartmann et al., 2002). Gibberellins can be applied successfully to break seed dormancy. Researchers (Gercekciolu and Cekic,
1999; Al Absi, 2010; Ghayyad et al., 2010) achieved highest seed
germination with GA3 1000 ppm application combined with stratication for 12 weeks at 24 Szab et al. (2012) reported that
the mean germination time of fresh harvested seed is decreased
by both gibberellin (GA4+7 and GA 3) treatments; the shortest was
when GA4+7 applied.0
One-year-old seedlings as liners are planted in the nursery
for budding. The rate of bud take on virus free, well established
seedlings is high and in the second year of nursery develop abundant branched one year old trees (Hrotk and Fzessry, 1996;
Stachowiak et al., 2014).
Softwood cuttings of P. mahaleb root easily, however the rooting of different clones vary from 50 to 100% (Sarger, 1972; Hrotk,
1982; Koleva, 2001; Ljubojevic et al., 2011; Szab et al., 2014). Szab
et al. (2014) improved the cutting production of stock-plants and
rooting of cuttings by biostimulator (Kelpak) application. Rooting
rate of hardwood cuttings is low, some years Bogdny showed
2030% rooting (Hrotk, unpublished data). Rooted softwood cuttings usually need one year more raising before lined up into
budding eld of the nursery, but using large cuttings (3035 cm)
usually results proper liners quality.
Rooting of microcuttings under in-vitro conditions is rather difcult, however, Giorgio and Standardi (1996), Muna et al. (1999),
Doric et al. (2014) reported about successful micro-propagation
results of P. mahaleb.

4.4. Training system and pruning

Mahaleb is the usual rootstock for Spanish Bush (SB) in dry
areas and on alkaline soil in Mediterranean countries (Spain, Italy,
Greece) (Negueroles, 2005; Iglesias and Peris, 2007), while Long
et al. (2015) suggest semi vigorous and vigorous rootstocks for
KGB, Steep leader, Tall spindle and UFO systems. Hrotk (2010)
reports on successful establishing of Hungarian Cherry Spindle,
when the soil fertility, rootstock vigor and cultivar considered
together combined with appropriate pruning protocol. Additional
root restriction by mechanical pruning could compensate the rootstock vigor and improve the precocity when applied in combination
on spindle trained trees under high density conditions (Hrotk,
2010). Among the cherry rootstocks the non-suckering mahaleb
root tolerate best the root pruning.

4.5. Environmental adaptability

Cold hardiness is an important attribute of rootstocks and rootstocks can also affect the response of the scion to cold temperatures
(Howell and Perry, 1990). Albeit P. cerasus and Prunus fruticosa are
considered the hardiest rootstocks, mahaleb is hardier than Mazzard. Within the mahaleb species the broad leaved subspecies is
hardier than the small leaved subspecies. Rootstock test in nursery
stage showed sensitivity of SL 64 to early frost injury. Extreme low
temperature (17 C) in middle of November resulted severe frost
damage on growing shoot system of SL 64, while the seedlings of
Korponay (broad-leaved) with terminated shoots were not injured
(Hrotk, 2004). P. avium is the least hardy species (Perry, 1987)
within Eucerasus, although Kppers (1978) reports some differences in hardiness of Mazzard selections.
Drought and heat tolerance of rootstocks is essential in many
cherry growing regions and this attribute may be linked to root
depth. Shallow-rooted dwarng rootstocks (some dwarng interspecic hybrids, P. cerasus and P. fruticosa) are more susceptible
to drought and heat injury. Due to their extensive root system the
most tolerant rootstocks appear to be the P. mahaleb selections and
hybrids (MxM series) (Perry, 1987). However, modern high density
orchards are usually irrigated, the drought and heat tolerance is still
an important rootstock trait in dry areas (Negueroles, 2005; Ercisli
et al., 2006; Iglesias and Peris, 2007; Hrotk, 2010). The deep penetrating roots of mahaleb cherry (Perry, 1987), the efcient water
uptake in root/soil interface, as well as the vessel transport capacity
in stem (Goncalves et al., 2007; Vgvri et al., 2008) and the higher
stomatal conductance of healthy leaf canopy on vigorous rootstocks
(Tombesi et al., 2010; Gyeviki et al., 2012) may contribute to the
higher drought tolerance of mahaleb rootstocks.
Adaptability to different soil conditions is considered as important rootstock trait. P. mahaleb and derivatives tolerate light sandy
and gravel soils with high lime content and pH levels of 7.88.5.
Mahaleb seedlings (Cema) proved to be tolerant to the calcareous and high pH soils in Shaanxi province of China, where in the
summer, during the rainy season, anaerobic conditions may cause
iron chlorosis when using P. pseudocerasus as rootstock (Faust et al.,
1998; Cai et al., 2007; Cai, 2015). Similarly on light sandy soil close
to Inner Mongolia mahaleb cherry as rootstock showed good adaptability.
In a grafted tree the rootstock is responsible for the uptake
of mineral nutrients, but can be selective in nutrient uptake and
transport, which results in different concentration of nutrients
reaching the leaves and fruits. However, literature data are inconsistent concerning the uptake efciency of different rootstocks
(Hanson and Proebsting, 1996; Sitarek et al., 1998; Usenik et al.,
2008; Roversi et al., 2008). Results of Moreno et al. (1996) suggest that soil/rootstock interaction and adaptability can provide
balanced nutritional status for the scion. On dry, gravel calcareous soil all rootstocks induced low leaf Fe concentrations, although
visual chlorosis symptoms were not observed. In Van cultivar,
Adara rootstock followed by CAB 6P and Gisela 5 showed the most
balanced nutritional values. On the other hand, SL 64 had leaf mineral element concentrations below the optimum, probably due to
the bad adaptation to heavy soil conditions. Results of Hrotk et al.
(2014) conrm the rootstock effect on nutrient supply, and that
those rootstocks provide balanced nutrient supply, which t well
into soil conditions. They found that the efciency of GiSelA 6 root
is emphasized in uptake and supply of leaves with nitrogen (N),
phosphorus (P), potassium (K), zinc (Zn), boron (B), and manganese
(Mn), but trees on this rootstock tend to develop calcium (Ca), magnesium (Mg), and copper (Cu) deciencies on light sandy soil. P.
mahaleb rootstocks on calcareous sandy soil are efcient supplier
of N, P, K, Ca, Mg, Fe, and Cu, but this root tends to develop Zn,
B, and Mn deciencies. Stachowiak et al. (2015) also conrmed the

K. Hrotk / Scientia Horticulturae 205 (2016) 7078

higher Fe supply on Mahaleb roots. These data suggest that nutrient

deciencies or excesses might be corrected by a judicious rootstock
selection.
Mahaleb cherry as rootstock show sensitivity, tolerance or resistance to various soil-born pests and diseases, however several
contradicting data are known in this concern. Several nematode
species attack the roots of cherry trees. Zepp and Szczygiel (1985)
found that Pratylenchus penetrans Cobb attacks P. mahaleb roots
more readily than Mazzard and P. cerasus. By Webster and Schmidt
(1996) Mazzard and P. cerasus are more tolerant than P. mahaleb to
Meloidogyne incognita (Kofoid and White) Chitwood. In contrary by
Hartmann et al. (2002) mahaleb roots are more resistant to rootlesion nematode (Pratylenchus vulnus) than Mazzard. They are also
resistant to the root-knot nematode (Meloidogyne incognita) but
susceptible to M. javanica.
On heavy soils with low drainage capacity or in waterlogging
sites Phytophthora species may cause serious tree decay on the susceptible P. mahaleb root, while P. cerasus and Mazzard are more
tolerant (Wicks et al., 1984; Cummins et al., 1986). MxM series
(Perry, 1987) and series of mahaleb cherry selected at Davis University of California (UCMH 55, 56, 59) tolerate Phytophtora infection
and can be recommended even on heavy soil. All rootstocks are
sensitive to Verticillium and there is no known source of resistance.
In the U.S.A. where Armillaria mellea can cause root damage, P.
mahaleb, P. cerasus, Colt and Inmil were found to be sensitive, Mazzard less sensitive, and MxM 60 showed the least sensitivity (Proffer
et al., 1988). Leaf spot caused by Blumeriella jaapii can cause severe
leaf fall in nursery liners. Only P. mahaleb is tolerant whereas P.
avium, P. cerasus and their derivatives are more or less sensitive
(Wharton et al. 2003; Schuster, 2004).
Bacterial diseases that create problems include crown gall
(Agrobacterium tumefaciens) which infects trees in the nursery
as well as in orchards where it can reduce growth and productivity. From among the cherry rootstocks less sensitive are the
mahaleb cherry rootstocks and P. fruticosa hybrids while Colt and
the Mazzard clone F12/1 are both sensitive. Pseudomonas s. pv.
mors-prunorum and Pseudomonas syringae pv. syringae or bacterial canker is a particularly damaging disease in the humid zone of
temperate areas. Mahalebs are known to be tolerant, whereas Mazzard genotypes are considered susceptible (Webster and Schmidt,
1996).
There is no known resistance to viruses or phytoplasmas,
although there are considerable differences in sensitivity. In France
P. mahaleb rootstocks are more susceptible to leafhopper transmitted phytoplasma (Moli<res decline) compared to trees on
Mazzard. Western X Disease in the U.S.A. which is also transmitted by leafhoppers can infect trees on Mazzard and Colt whereas
P. mahaleb rootstocks are hypersensitive. Albeit mahaleb, MxM 2
and MxM 46 are considered to be resistant, but in a grafted
tree when the phytoplasma reaches the graft union by blockage
of its movement from the scion into the rootstock, causes rapid
scion death (Beckman and Lang, 2003). Mild strains of two common ilarviruses, prunus necrotic ringspot (PNRSV) and prune dwarf
(PDV) may cause similar symptoms on sensitive rootstocks but
result in little or no negative impact for cherries grown on mazzard
or mahaleb rootstocks.

5. Achievements of P. mahaleb breeding as cherry rootstock

5.1. Seed tree selection and establishing of seed orchards
Seed orchards with selected seed trees are planted to provide
the nursery industry with a regular supply of high quality seeds
that are either hybrid seeds or inbred lines. The early seed orchards
which were initially selected from wild populations based on the

75

phenotype characteristics (e.g. healthy, vigorous tree, regular seed

crop) now have given way to evaluation of the progeny of the seed
2006). The ower
trees from known pollinations (Hrotk and Erdos,
fertility of these genotypes determines the mating options within
the orchard and accordingly the genetic composition of the seedling
progeny. The owers in the majority of seed producing clones for
successful seed production need cross pollination, therefore the
seedling progeny is a hybrid with all attributes of hybrid vigor and
greater homogeneity in the phenotype of the F1 population than
former seed sources. Such F1 populations are produced for rootstock use (Kppers, 1978; Nyjt, 1987). Several seed orchards with
cross pollination can consist of three to ve clones, each pollinating the other (Funk, 1969; Nyjt, 1987; Perry, 1987). In this type
of hybrid mating system the progeny represents a hybrid family
of different mating combinations. Progenies of these seed orchards
are usually evaluated for their nursery and orchard value.
Seed sourced/mother trees selected for superior phenotypic
traits have been released from several countries (Table 1). Advantages of seed orchards include the potential for a virus-free seed
source, higher germination capacity, hybrid seed of known parents
and improved uniformity of orchard trees compared to openpollinated seeds (Kppers, 1978; Yoltuchovski, 1977; Nyjt, 1987;
2006).
Webster and Schmidt, 1996; Hrotk and Erdos,
Among the mahaleb genotypes some are self-fertile, like
Heimann X (Germany), Korponay (Hungary) and SL 405 (France),
their seedlings are slightly less vigorous (525%). Although the trees
are smaller, the seedlings juvenility could not be overcome by this
breeding method, the trees budded on them are not expected to
be precocious enough, thus their use can be recommended mainly
for mechanical harvested orchards and training systems requiring
semi vigorous or standard rootstocks (Hrotk, 2010; Long et al.,
2015).

5.2. Inbreeding
(1968), Fischer (1985), Claverie (1996),
Heimann (1932), Sebok
Hrotk (1996, 2004) and Hrotk and Magyar (1998) reported on
self-fertile types of P. mahaleb. Self-fertile seed trees may produce
a diversity in seedling characters and segregation in the population
(Hrotk and Magyar, 1998; Hrotk, 2004), which is more or less
tolerable among the I1 seedlings used for rootstocks. Among Korponay I1 seedlings occurred genotypes with yellow, red or black
fruit colour, various growth habit (e.g. columnar type), and various
level of ower fertility. Seedlings of the self-fertile genotype
of P. mahaleb Heimann X was known for having very uniform
progeny (Heimann, 1932; Kppers, 1978). In France, Claverie
(1996) and in Hungary Hrotk and Magyar (1998) reported on
utilization of inbreeding of P. mahaleb seed tree selection with
the aim of producing less vigorous and more uniform seedling
populations. No progress in tree size reduction and yield efciency
of Schattenmorelle trees could be achieved by Fischer (1985) on
inbred seedling lines. In contrary Hrotk et al. (2005) reported
sour cherry trees grafted
50% growth reduction on rdi btermo
on I2 inbred lines. The inbreeding of self-fertile seed trees could
provide a useful tool for rootstock breeding. Within inbred lines
Hrotk and Magyar (1998) and Hrotk (2004) reported on genetic
dwarf seedlings in the Heimann X progeny produced in pots
from all germinated seeds, which in a seedbed population might
have always been eliminated or sorted out. Jacob (1992) showed
similar genetic dwarf mahaleb trees in their breeding garden
in Geisenheim. Despite these opportunities no clonal selection
among inbred populations has been reported.

76

K. Hrotk / Scientia Horticulturae 205 (2016) 7078

5.3. Clonal rootstock selection

For the selection of clonal mahaleb cherry rootstocks, wild populations as well as seedlings produced for rootstocks in the nursery
provide appropriate genetic diversity. Major selection criteria
were: ease of vegetative propagation, cold hardiness, adaptability to different soil and climatic conditions, tolerance or resistance
to pests and diseases, freedom from suckering, graft compatibility,
tree longevity, tree size control, effect on scion precocity, increased
productivity, and fruit quality.
Due to the development of the propagation technology in the
last fty years several clonal rootstocks were selected from the
P. mahaleb. The achievements of these selection was the uniform
plant material for the nurseries and fruit growers, but considering
the vigor and precocity of trees on these rootstocks little progress
was made (Webster and Schmidt, 1996). The vegetative propagated
clonal P. mahaleb rootstocks show extended vigor range, some
of them are semi-vigorous with certain precocity. Genetic dwarf
genotypes also occurred in breeding projects, but they were still
not utilized as rootstock or in rootstock breeding.
The rst P. mahaleb clonal rootstock, Sainte Lucie 64 (SL 64),
(Table 2), was selected in France for its ease of propagation, compatibility with sweet cherries, and productivity in orchard conditions.
In the second half of the last century no more clonal mahaleb
rootstocks got into the commercial propagation, albeit further
clonal rootstocks are reported from Germany (Baumann, 1977),
from Italy (Giorgio and Standardi, 1996), from Turkey (Misirli et al.,
1996), and from Hungary (Hrotk, 2004) involving semi-vigorous
rootstocks too.
The progress made in the eld of training systems of pedestrian
orchards and the accumulated experiences with vigorous rootstocks (Iglesias and Peris, 2008; Hrotk 2010; Long et al., 2015) may
give a new prospect for semi-vigorous or vigorous clonal mahaleb
selections in those conditions, where tolerance to drought, hot
summer, poor soils, high pH and lime tolerance are such rootstock traits that can improve the cherry growing adaptability. There
are promising reports from Iran (Ganji and Khalighi 2006), from
Turkey Koc and Bilgener (2013) and from Serbia (Barac et al., 2014)
on variability of clonal mahaleb. New clonal mahaleb rootstocks
are registered and/or patented in the USA (Lang 2006), in Bulgaria
(Koleva, 2001; Sotirov, 2005, 2012, 2015; Sotirov et al., 2015) and
in Hungary (Table 2).

5.4. Interspecic hybrids of P. mahaleb

Successful hybridisation of P. mahaleb with P. avium was carried
out in Oregon (Westwood, 1978; Perry, 1987) resulting in the MxM
series and OCR 2, 3. Two of them (MxM 14 and MxM 97) are moderate vigorous, in certain sites on mahaleb-soil both are considered
as possible rootstock for semi-intensive or intensive orchards. Their
resistance to Phytophtora, moderate vigor (MxM 14 and MxM 97)
and improved precocity contributed to wider application in USA
and Southern and Central European countries.
Crosses between P. mahaleb and P. fruticosa have been reported
by De Palma et al. (1996) and Hrotk (2004); testing is still in early
stages. The creation of hybrids, huge efforts were made to overcome low ower fertility (De Palma et al., 1996). In Central and
East-Europe the native geographic areal of P. mahaleb overlaps with
P. avium and P. fruitcosa, which in certain years allows spontaneous
crosses (Krpti, 1944; Wojcicki, 1991; Hrotk and Facsar, 1996).
Spontaneous hybrids of P. mahaleb and P. fruticosa have been collected in Hungary (Hrotk and Facsar 1996; Hrotk 2004); testing is
in early stages. On every location P. fruticosa f. aucta Borb. was found
as a third hybrid group. Some individuals of this hybrid group could
be clearly identied as Prunus x javorkae Krp. (=Prunus fruticosa

P. mahaleb). In the ambient forest (thickets) there are many trees

of P. mahaleb L., thus the conditions for the crossing were given.
Recently Lichev et al. (2014) evaluated hybrid rootstock
between P. mahaleb and GiSelA 5 (Hybrid 2/10) as interstock in
early stage of trees. Such mahaleb crosses with dwarf and precocious rootstocks may promise the desired progress in rootstock
breeding.

6. Conclusion
Adaptability of mahaleb cherry to continental climate, tolerance
to drought, hot summer, poor soils, high pH and lime tolerance
are important rootstock traits that can improve the cherry growing adaptability under those environmental conditions that due
to the climate changes are usually forecasted. Albeit some of the
modern orchard systems for pedestrian orchards prefer dwarng
and precocious rootstocks, the diverse rootstock usage can be forecasted in sweet and sour cherry growing, when training system
and rootstock are considered together, matched properly with the
vigor of orchard site. Traditional seed tree selection resulted in seed
tree clone, producing high quality seedlings for sweet- and sour
cherry growing but their vigor control and precocity does not meet
all the requirements of modern pedestrian orchards. Variability of
mahaleb cherry manifested in selected genotypes, inbred lines and
potential of interspecic crosses promise progress in those directions that are desired rootstock traits in vigor control, precocity
and environmental adaptability. Further breeding could help with
selection spur type clones, from among crosses with genetic dwarf
types or inbred lines. Interspecic hybrids with P. avium proved to
be useful, further crosses with P. fruticosa and hybrid dwarf and
precocious rootstocks may be of interest.

References
Al Absi, K.M., 2010. The effects of different pre-sowing seed treatments on
breaking the dormancy of mahaleb cherries, Prunus mahaleb L. seeds. Seed Sci.
Technol. 38, 332340.
Balmer, M., Blanke, M., 2005. Developments in high density cherries in Germany.
Acta Hortic. 667, 273278.
Balmer, M., 2008. Evaluation of semi-dwarng rootstocks for sweet cherry
orchards in the Rhine river valley (Germany). Acta Hortic. 795, 203208.
G., Ognjanov, V., Obreht, D., Ljubojevic,
M., Bosnjakovic,
D., Pejic,
I., Gasic,
K.,
Barac,
2014. Genotypic and phenotypic diversity of cherry species collected in Serbia.
Plant Mol. Biol. Rep. 32 (1), 92108.
Barya, P., Kapan, M., 2005. The estimation of the growth and the branching of the
six stocks under the cherry and the sweet cherry trees. Acta Sci. Pol. Hortcult. 4
(1), 119129.
Baumann, G., 1977. Clonal selection in Prunus mahaleb rootstocks. Acta Hortic. 75,
139148.
Beckman, T.G., Lang, G.A., 2003. Rootstock breeding for stone fruits. Acta Hortic.
622, 531551.
Bujdos, G., Hrotk, K., 2014. Preliminary results on growth: yield and fruit size of
some new precocious sweet cherry cultivars on Hungarian bred mahaleb
rootstocks. Acta Hortic. 1058, 559564.
Cai, Y.L., Cao, D.W., Zhao, G.F., 2007. Studies on genetic variation in cherry
germplasm using RAPD analysis. Sci. Hort. 111, 248254.
Cai, Y. 2015. Personal communication.
Cittadini, E.D., van Keulen, H., Peri, P.L., Ridder, N., 2007. Designing a target-tree for
maximizing gross value of product in patagonian sweet cherry orchards. Int. J.
Fruit Sci. 6, 322.
Claverie, J., 1996. New selections and approaches for the development of cherry
rootstocks in France. Acta Hortic., 373376.
Cummins, J.N., Wilcox, W.F., Forsline, P.L., 1986. Tolerance of some new cherry
roootstocks to December freezing and to Phytophtora root rots. Compact Fruit
Tree 19, 9093.
De Palma, L., Palasciano, M., Godini, A., 1996. Interspecic hybridization program
aimed at obtaining dwarng and non-suckering rootstocks for sweet cherry.
Acta Hortic. 410, 177181.
De Salvador, F.R., Di Tomaso, G., Bonoglio, P., Piccioni, C., 2005. Performance of
new and standard cherry rootstocks in different soils and climatic conditions.
Acta Hortic. 667, 191200.
D., Ognjanov, V., Ljubojevic Barac,
M.G., Dulic,
J., Pranjic,
A., Dugalic,
K., 2014.
Doric,
Rapid propagation of sweet and sour cherry rootstocks. Not. Bot. Horti
Agrobot. 42 (2), 488494.

K. Hrotk / Scientia Horticulturae 205 (2016) 7078

Edin, M., Garcin, A., Lichou, J., Jourdain, J.M., 1996. Inuence of dwarng cherry
rootstocks on fruit production. Acta Hortic. 410, 239243.
Ercisli, S, Esitken, A, Orhan, E, Ozdemir, O. 2006. Rootstocks used for temperate
fruit trees in turkey: an overview Scientic Works of The Lithuanian Institute
of Horticulture and Lithuanian University of Agriculture. Sodininkyste Ir
Darzininkyste 25: 2733.
Faust, M., Surnyi, D., 1997. Origin and dissemination of cherry. Hortic. Rev. 19,
263317.
Faust, M., Deng, X., Hrotk, K., 1998. Development project for cherry growing in
Shaanxi province of China P. R. Acta Hortic. 2 (468), 763769.
Faust, M., 1989. Physiology of Temperate Zone Fruit Trees. John Wiley and Sons
Inc., New York, Singapore.
Favr-Bonvin, J., Massias, M., Mentzer, C., Massicot, J., 1968. Siles de biosynthese
des coumarines chez Prunus Mahaleb. Phytochemistry 7, 15551560.
Feucht, W., Vogel, T., Scimmelpfeng, H., Treutter, D., Zinkernagel, V., 2001.
Kirschen-und Zwetschenanbau. E. Ulmer GmbH Co., Stuttgart.
Feucht, W., 1982. Das Obstgehlz. Eugen Ulmer Verlag, Stuttgart.
Fischer, M., 1985. Selektionsarbeiten an Prunus mahaleb L. als Unterlagen fr Sund Sauerkirschen. Arch. Gartenbau 33 (2), 75.
Funk, T., 1969. Bericht ber die Selektion bei Prunus mahaleb L. in der Deutschen
Demokratischen Republik. Arch. Gartenbau 4, 219237.
Ganji, M.E., Khalighi, A., 2006. Genetic variation of mahaleb (Prunus mahaleb L.) on
some iranian populations using morphological characters. J. Appl. Sci. 6 (3),
651653.
Garner, R., 1967. The cherry rootstock position today. An.Rep.East Mal.Res.Sta.
Maidstone/Kent, 212214.
Gercekciolu, R., Cekic, C., 1999. The effects of some treatments on germination of
mahaleb (Prunus mahaleb L.) seeds. Turk. J. Agric. For. 23, 145150.
Ghayyad, M., Kurbysa, M., Napolsy, G., 2010. Effect of endocarp removal,
gibberelline, stratication and sulfuric acid on germination of mahaleb (Prunus
mahaleb L.) seeds. American-Eurasian. J. Agric. Environ. Sci. 9, 163168, ISSN
1818-6769.
Giorgio, V., Standardi, A., 1996. Growth and production of two sweet cherry
cultivars grafted on 60 ecotypes of Prunus mahaleb. Acta Hortic. 410, 471475.
Goncalves, B., Correia, C.M., Silva, A.P., Bacelar, E.A., Santos, A., Ferreira, H.,
Moutinho-Pereira, J.M., 2007. Variation in xylem structure and function in
roots and stems of scionrootstock combinations of sweet cherry tree (Prunus
avium L.). Trees 21 (2), 121130.
Gyeviki, M., Hrotk, K., Hon, P., 2012. Comparison of leaf population of sweet
cherry (Prunus avium L.) trees on different rootstocks. Sci. Hortic. 141, 3036.
Hartmann, H.T., Kester, D.E., Davies, F.T., Geneve, R.L., 2002. Plant Propagation.
Prentice-Hall, Inc., USA, New Jersey, pp. 770.
Heimann, O.R., 1932. Zur Frage der Selektion der Steinweichsel Prunus
mahalebPrunus mahaleb als Veredlungsunterlage fr Virschen. Der Obst und
Gemsebau Bln. 78, 138.
Howell, G.S., Perry, R.L., 1990. Inuence of cherry rootstocks on the cold hardiness
of twigs of the cherry scion cultivar. Sci. Hortic. 43, 103108.
Hrotk, K., 2004. Cherry rootstock breeding at the department of Fruit Science,
Budapest. Acta Hortic. 658, 491495.
Hrotk, K., Cai, Y.L., 2014. Development in intensive cherry orchard systems in
Hungary and China. COST FA 1104 Meeting: Sustainable production of
high-quality cherries. Bordeaux, 1315. October 2014. Book of Abstracts, page
10.
Hrotk, K., Erd"os, Z., 2006. Floral biology of tree fruit rootstocks. Int. J. Hortic. Sci.
12 (2), 153161.
Hrotk, K., Fzessry, A., 1996. Effect of rootstocks on the branching and quality of
cherry trees in the nursery. Acta Hortic. 410, 507510.
Hrotk, K., Facsar, G., 1996. Taxonomic classication of Hungarian populations of
Prunus fructicosa (Pall.) Woronow hybrids. Acta Hortic. 410, 495498.
Hrotk, K., Magyar, L., 1998. Inbreeding of Prunus mahaleb. Acta Hortic. 468,
393400.
Hrotk, K., Magyar, L., 2004. Rootstocks for cherries from department of fruit
science, Budapest. Int. J. Hortic. Sci. 10 (3), 6366.
Hrotk, K., Magyar, L., Simon, G., 1999. Growth and yield of sweet cherry trees on
different rootstocks. Int. J. Hortic. Sci. 5, 98101.
Hrotk, K., Magyar, L., Szab, S., 2005. Growth and productivity of rdi b"oterm"o
sour cherry trees on mahaleb inbred lines. Acta Hortic. 667, 261268.
Hrotk, K., Magyar, L., Gyeviki, M., 2008. Evaluation of native hybrids of Prunus
fruticosa pall. As cherry interstocks. Acta Agr. Serbica 13 (25), 4145.
Hrotk, K., Magyar, L., Gyeviki, M., 2009a. Effect of rootstocks on vigor and
productivity in high density cherry orchards. Acta Hortic. 825, 245250.
Hrotk, K., Magyar, L., Gyeviki, M., 2009b. Effect of rootstoks on growth and yield
of Carmen sweet cherry trees. Bull. UASVM Horticul. 66 (1), 143148.
Hrotk, K., Magyar, L., Hoffmann, S., Gyeviki, M., 2009c. Rootstock evaluation in
intensive sweet cherry (Prunus avium L.) orchard. Int. J. Hortic. Sci. 15 (3), 712.
Hrotk, K., Magyar, L., Borsos, G., Gyeviki, M, 2014. Rootstock effect on nutrient
concentration of sweet cherry leaves. J. Plant Nutr. 37 (9), 13951409.
Hrotk, K., 1982. Sajmeggy alanyklnok szaportsa zlddugvnyozssal
(Propagation of Mahaleb cherry by softwood cuttings). Kertgazdasg 4, 4550.
Hrotk, K., 1990. The effect of rootstocks on the growth and yield of Meteor korai
sour cherry variety, XXIII. Int. Horticult. Congress, Abstracts 2., Poster Nr. 4165.
Hrotk, K., 1996. Variability in Prunus mahaleb L. for rootstock breeding. Acta
Hortic. 410, 183188.
Hrotk, K., 2008. Progress in cherry rootstock research. Acta Hortic. 795, 171178.
Hrotk, K., 2010. Intensive cherry orchard systems and rootstocks from Hungary.
Compact Fruit Tree 43 (1), 510.

77

Ieri, F., Pinelli, P., Romani, A., 2012. Simultaneous determination of anthocyanins,
coumarins and phenolic acids in fruits, kernels and liqueur of Prunus mahaleb
L. Food Chem. 135 (4), 21572162.
Iglesias, I., Peris, M., 2008. Speciale Ciliego. La produzione spagnola vince grazie a
precocit, qualit e organizzazione tecnico-commerciale. Frutticoltura 3,
2026.
Jacob, H., 1992. Personal communication.
Krpti, Z., 1944. Vizsglatok a hazai Cerasus alnemzetsgbe tartoz hazai
Prunusokon, 10. Bulletin of the Hungarian College for Horticulture and
Vineculture, Budapest, pp. 6680.
Kppers, H., 1978. Problematik der Veredlungsunterlagen fr Sauer- und
Skirschen im Spiegel von 250 Jahren. Dt. Baumsch. 11. 350359.
Kappel, F., Lang, G., Anderson, L., Azarenko, A., Facteau, T., Gaus, A., Southwick, S.,
2005. NC-140 regional cherry rootstock trial (1998): results from western
North America. Acta Hortic. 667, 223232.
Khadivi-Khub, A., Zamani, Z., Fatahi, M.R., 2012. Multivariate analysis of Prunus
subgen. Cerasus germplasm in Iran using morphological variables. Genet.
Resour. Crop Evol. 59 (5), 909926.
Koc, A., Bilgener, S., 2013. Morphological characterization of cherry rootstock
candidates selected from Samsun Province in Turkey Turk. J. Agric. For. 37,
110.
Koleva, A., 2001. Rootstocks. In: Borovinova, Georgiev V.M., Koleva, A. (Eds.), Sweet
Cherry, Zemizdat. EOOD, Soa, pp. 186207.
Krssmann, G., 1978a. Die Baumschule. Verlag Paul Parey, Berlin-Hamburg.
Krssmann, G. 1978. Handbuch der Laubgehlze. 2. Au. Bd. III. Verlag Paul Parey.
Lang, G., 2000. Precocious, dwarng, and productivehow will new cherry
rootstocks impact the sweet cherry industry? HortTechnology 10 (4), 719725.
Lang, G. 2006 Cherry rootstocks in Clark, in: J.R. Finn, C.E. (Eds). Register of New
Fruit and Nut Cultivars List 43. HortScience, 41 (5): 11091110.
Lang, G.A. Producing rst class sweet cherries: integrating new technologies,
germplasm and physiology into innovative orchard management. In
Proceedings of the 3rd Conference Innovation in Fruit Growing. Belgrade,
2011, 59-74.
Lichev, V., Papachatzis, A., 2009. Results from the 11-year evaluation of 10
rootstocks of the sweet cherry cultivar stella. Acta Hortic. 825, 513520.
Lichev, V., Botu, M., Papachatzis, A., 2014. First results from the examination of
three interstocks for the sweet cherry cultivar Stella. Acta Hortic. 1020,
381384.
M., Osterc, G., Ognjanov, V., Barac, G., Bosnjakovic, D., Mladenovic, E.,
Ljubojevic,
Cukanovic, J. 2011. Umnozavanje selekcija slabobujnih vegetativnih podloga za

2011, vol. 45, br. 173174, str.

visnju i tresnju zelenim reznicama. Vocarstvo,
4954.

R., Cahn, H., 2005. Developments in high density

Long, L., Facteau, T., Nunez-Elisea,
cherries in the USA. Acta Hortic. 667, 303310.
Long, L., Lang, G., Musacchi, S. and Whiting, M. 2015. Cherry training systems. PNW
667. Oregon State University Extension Service pp. 63.
Magyar, L., Hrotk, K., 2013. The effect of rootstock and spacing on the growth and
yield of Kntorjnosi sour cherry variety in intensive orchard. Acta Hortic.
981, 373378.
T., Milosevic,
N., Milivojevic,
J., Glisic,
I., Nikolic,
R., 2014. Experiences
Milosevic,
with Mazzard and Colt sweet cherry rootstocks in Serbia which are used for
high density planting system under heavy and acidic soil conditions. Sci.
Hortic. 176, 261272.
Misirli, A., Glcan, R., Tanrisever, A., 1996. The relation between tree vigour of
Prunus mahaleb L. types and sieve tube size in phloem tissue. Acta Hortic. 410,
227232.
Moreno, M.A., Montanes, L., Tabuenca, M.C., Cambra, R., 1996. The performance of
Adara as a cherry rootstock. Sci. Hortic. 65 (1), 8591.
Muna, A.S., Ahmad, A.K., Mahmoud, K., Abdul-Rahman, K., 1999. In vitro
propagation of a semi-dwarng cherry rootstock. Plant Cell. Tissue Organ Cult.
59, 203208.
Musacchi, S., Gagliardi, F., Serra, S., 2015. New training systems for high-density
planting of sweet cherry. HortScience 50 (1), 5967.
Negueroles, P.J., 2005. Cherry cultivation in Spain. Acta Hortic. 2 (667), 293301.
Nyjt, F., 1987. Az alanykutats hazai eredmnyei. Kertgazdasg 19 (5), 934.
Ozturk, I., Karaman, S., Baslar, M., Cam, M., Caliskan, O., Sagdic, O., Yalcin, H., 2014.
Aroma, sugar and anthocyanin prole of fruit and seed of mahlab (Prunus
mahaleb L.): Optimization of bioactive compounds extraction by simplex
lattice mixture design. Food Anal. Methods 7 (4), 761773.
Perry, R.L., 1987. Cherry rootstocks. In: Rom, R.C., Carlson, R.F. (Eds.), Rootstocks for
Fruit Crops. John Wiley & Sons, New York, pp. 217264.
Proffer, T.J., Jones, A.L., Perry, R.L., 1988. Testing of cherry rootstocks for resistance
to infection by species of Armillaria. Plant Dis. 72, 488490.
Rehder, A., 1990. Manual of Cultivated Trees and Shrubshardy in North America.
Dioscorides Press, Portland, Oregon, US, pp. 996.
Rejman, A., Scibisz, K., Czarnecki, B., 2002. Szkkarstwo roslin sadowniczych.
PWRiL, Warszawa.
Robinson, T.L., 2005. Developments in high density sweet cherry pruning and
training system around the world. Acta Hortic. 2 (667), 269272.
Roversi, A., Ughini, V., Monteforte, A., 2008. Inuence of genotype, year and soil
composition on sweet cherry mineral composition. Acta Hortic. 795, 739745.
Sansavini, S., Lugli, S., 2014. New rootstocks for intensive sweet cherry plantations.
Acta Hortic. 1020, 411434.
Sarger, J., 1972. Le bouturage ligneux de lespece Prunus mahaleb , Convegno del
ciliego, Verona, 1972 June.

78

K. Hrotk / Scientia Horticulturae 205 (2016) 7078

Saunier, R., Claverie, J., 2001. Le cerisier: volution de la culture en France et dans
le monde Point sur les varits, les portegreffe. La Fruit Belge 490, 5062.
Schnberg, G., 1963. Gewinnung vegetativ vermehrter Unterlagen von Prunus
mahaleb. Obstbau 6, 9092.
Schuster, M., 2004. Investigation on resistance to leaf spot disease, Blumeriella
jaapii in cherries. J. Fruit Ornam. Plant Res. 12, 275279.
Seb"ok, L., 1968. Sajmeggy magfk szelekcija (Selection of mahaleb cherry seed
trees.) Sz"ol"o- s Gymlcstermeszts. Budapest 4, 133143.
Shahi-Gharahlar, A., Zamani, Z., Fatahi, R., Bouzari, N., 2011. Estimation of genetic
diversity in some Iranian wild Prunus subgenus Cerasus accessions using
inter-simple sequence repeat (ISSR) markers. Biochem. Syst. Ecol. 39 (4),
826833.
Simon, G., Hrotk, K., Magyar, L., 2004. Fruit quality of sweet cherry cultivars
grafted on four different rootstocks. Int. J. Hortic. Sci. 10, 5962.
Sitarek, M., Gryzb, Z.S., Olszewski, T., 1998. The mineral elements concentration in
leaves of two sweet cherry cultivars grafted on different rootstocks. Acta
Hortic. 468, 373376.
Sotirov, D., Antonova, V., Hrisztov, N., Krumov, S., Stoianov, H., 2015.
Manifestations f Summit sweet cherry cultivar on different interstems. Plant
Sci. (Bulgaria) 52 (2), 37.
Sotirov, D., 2005. Growth and Reproductive Characteristics of Sour Cherry Cultivars
Grown on Own Roots and Grafted on IK-M9 Mahaleb Rootstock, vol. 3.
Scientic Works of National Center for Agrarian Sciences, Soa, pp. 6771.
Sotirov, D., 2012. Growth characteristics of van sweet cherry cultivar grafted on six
rootstocks. Plant Sci. (Bulgaria) 49, 5560.
Sotirov, D.K., 2015. Performance of sweet cherry cultivars Van and Kozerska on
clonal rootstocks. Acta Hortic. 1099, 727733.
Stachowiak, A., Swierczynski, S., Kolasinski, M., 2014. Growth and yielding of
sweet cherry trees grafted on new biotypes of Prunus mahaleb (L.). Acta Sci. Pol.
Hort. Cult. 13 (5).

nski,
S., Kolasinski,
M., 2015. Inuence of
Stachowiak, A., Bosiacki, M., Swierczy
rootstocks on different sweet cherry cultivars and accumulation of heavy
metals in leaves and fruit. Horticul. Sci. (Prague) 42 (4), 193202.
Szab, V., Magyar, L., Mndy, A., Hrotk, K., 2012. Germination of Prunus mahaleb L:
seeds by gibberellic acid (GA) treatments in different seed age. Eur. J. Hortic.
Sci. 77 (5), 199203.
Szab, V., Nmeth, Zs., Srvri, A., Vgvri, Gy., Hrotk, K., 2014. Effects of
biostimulator and leaf fertilizers on Prunus mahaleb L. stockplants and their
cuttings. Acta Sci. Pol. Hort. Cult. 13 (6), 113125.
Tanrisever, A., Feucht, W., 1978. Gehalt an Flavanen im Phloem von Kirschartigen
Prunusgehlzen. Mitt. Klosterneuburg 28, 7274.

Tatarinov, A.N., Zuev, V.F., 1984. Pitomnik plodovyh y yagodnych kultur.

Rosselhosisdat, Moscow.
Terp, A., 1968. A sajmeggy (Cerasus mahaleb (L.) Mill.) taxonomiai problmi s a
gyakorlat. (Problems of taxonomy of Cerasus mahaleb (L.) Mill.)., Szl- s
Gy"omlcstermeszts. Budapest 4, 103131.
Thomas, M. s Sarger, J. 1965. Selection de Prunus mahaleb porte greffe du cerisier,
Rapport general du Congress Pomologique de Bordeaux, 175201.
Tombesi, S., Johnson, R.S., Day, K.R., DeJong, T.M., 2010. Relationships between
xylem vessel characteristics, calculated axial hydraulic conductance and
size-controlling capacity of peach rootstocks. Ann. Bot. 105 (2), 327331.
Treutter, D., Feucht, W., Liebster, G., 1993. 40 Jahre Wissenschaft fr Den Obstbau
in Weihenstephan. Obst- und Gartenbauverlag, Mchen, pp. 170.
Usenik, V., Stampar, F., Fajt, N., 2008. Sweet cherry rootstock testing in Slovenia.
Acta Hortic., 273276.
Vgvri, G., Hrotk, K., Magyar, L., Hajagos, A., Csigai, K., 2008. Histological
investigation of cherry rootstocks. Acta Hortic. 795, 339344.
Webster, A.D., Schmidt, H., 1996. Rootstocks for sweet and sour cherries. In:
Webster, A.D., Looney, N.E. (Eds.), Cherries: Crop Physiology, Production and
Uses. CAB International, Wallingford, UK, pp. 127163.
Webster, A.D., 1998. Strategies for controlling the tree size of sweet cherry trees.
Acta Hortic. 410, 229240.
Westwood, M.N., 1978. Mahaleb Mazzard Hybrid Cherry Stocks. Fruit Var. J., pp.
3239.
Wharton, P.S., Iezzoni, A., Jones, A.L., 2003. Screening cherry germ plasm for
resistance to leaf spot. Plant Dis. 87, 471477.
Wicks, t, J., Bumbieris, M., Warcup, J.H., Wallace, HR, 1984. Phytophtora in Fruit
Orchards in South Australia. Biennial Rep. of the Waite Agricultural Research
institute, pp. 147.
Wojcicki, J.J., 1991. Prunus stacei (Rosaceae), a new spontaneous hybrid of P.
fruticosa, P. cerasus and P. avium. Fragm. Flor. Geobot. 35 (12), 14139.
Yoltuchovski, M.K., 1977. Proisvodstvenno-biologicheskaya characteristika
raslychnych form podvoyev vishnyi. In plodovoje pitomnikovodstvo Moldavii.
Redact. Andryushtchenko. Isdatelstvo Krtya Moldavenyanske. Kishinau, 3969.
Zepp, L., Szczygiel, A., 1985. Pathogenicity of Pratylenchus crenatus and
Pratylenchus neglectus to three fruit tree seedling rootstocks. Fruit Sci. Rep. 12
(3), 109117.
de Haas, P.G., Hildebrandt, W., 1967. Die Unterlagen und Baumformen des Kernund Steinobstes. Ulmer Verlag, Stuttgart.