Cultivated black-fruited aronia:
place, time, and probable mechanism of formation
A.K. Skvortsov, Yu.K. Maitulina, Yu.N. Gorbunov
Bull. MOIP, Otd. Biol. 1983, 88(3): 88-96
Translation: Irina Kadis
The black-fruited aronia, which is so widespread in cultivation in the
USSR, is devoid of any reliably detectable variability, at the same time
differing significantly from the wild, American black-fruited aronia. This
may provide grounds for recognizing it as a distinct species, Aronia
mitschurinii Skvortsov et Maitulina (Skvortsov, Maitulina 1982).
In this communication, we are attempting to clarify the time, place,
and biological mechanism of cultivated aronia formation. For this purpose we
have reconstructed to the best of our ability the history of its
introduction in this country and also studied its flowering, fruiting, and
I. History of Introduction
The first reference to the presence of black-fruited aronia in Russia
is found in the catalog of plants published by the Kremenets Botanical
Garden, now in Ternopil Oblast of West Ukraine, under the name of
Mespilus melanocarpa (Besser 1816).
In 1823, M. melanocarpa was already
cultivated in the Kharkov University Botanical Garden, Ukraine (Index
plantarum... 1823). In 1831 it was grown in the Kwitka[-Osnovyanenko] Park,
Moskalyovka District, Kharkov (Catalogue des plantes ... 1831). A
herbarium sample collected in 1825 on the Crimea Peninsula, the Black Sea
Coast, Ukraine and labeled M. melanocarpa is preserved in the Moscow
University Herbarium. Morphologically this sample is no different from North
At about the same time black-fruited aronia arrives to St. Petersburg
and Moscow. M. melanocarpa was listed in 1824 catalog of plants grown in the
St. Petersburg Botanic Garden (Index plantarum ... 1824) and the Emperor's
Garden in Pavlovsk, a suburb of St. Petersburg (Weinmann 1824, as Pyrus
melanocarpa Willd.). In 1825 the plant makes it to the Index seminum of the
Moscow University Botanic Garden (Index seminum ... 1825).
The first brief description of the black-fruited aronia in Russian was
published by F.B. Fischer in 1839. He noted that the plant could endure the
northern climate and would beautify any nursery (1839, vol. 2: 181). Later
on, E.L. Regel (1874), R.I. Schreder (1899), and E.L. Wolf (1915) also
mentioned aronia's hardiness sufficient for it to exist in St. Petersburg
Early in the 20th century black-fruited aronia was introduced to the
Caucasus. Seed originating from the Caucasus botanic gardens were listed in
the 1909 exchange list of the St. Petersburg Botanic Garden (Delectus
seminum ... 1909). However, it never became widespread in the Caucasus and
apparently soon disappeared, as we never succeeded finding any later
reference to its presence there.
Early 20th century was the time of the introduction of black-fruited
aronia to Siberia. According to 1913 Delectus seminum of the St. Petersburg
Botanic Garden, it was growing in Pavel S. Komissarov's garden in a suburb
Pavel Savvich Komissarov, Michurin's contemporary, is considered
the founder of the Siberian horticulture; his former garden is now one of
the state reservations.
See more at http://www.omskmap.ru/point/achair/person/72
Before the 20th century aronia was cultivated exclusively as an
ornamental plant. In all descriptions authors would refer to ornamental
qualities of its flowers and shiny black fruit. Any notes on the taste of
the fruit are absent from descriptions up until the early 20th century. It
was I.V. Michurin (Mitschurin) who first attempted growing black-fruited
aronia as an edible plant in the 90's of the 19th century. In his nursery in
Kozlov (now Michurinsk), he had a few bushes of black-fruited aronia
originating from Germany (Michurin 1948a). Exactly what propagules were
received by Michurin remains unknown. Some (Barabash 1960) believe it was
seed; others (Protsenko 1971, Kalinina 1973) write about cuttings. It is
also unknown whether the received material already constituted a new
large-fruited form or if this form was somehow first developed by
To our opinion, there is enough evidence to believe that the latter is
true. In his notebook (1948b: 345), among other new cultivars, Michurin
lists "a mountain-ash ... (omission) received through selection from three
generations of an acclimated variety." There was a total of four species of
introduced mountain-ashes in Michurin's garden: Sorbus aria, S. alpina,
S. latifolia, and S. melanocarpa. Since he never recommended the first three
species for cultivation, we have to conclude that the note spoke about a new
form of black-fruited aronia. Additionally, in another place he wrote: " I
have introduced a few improved kinds of fruiting plants, among which ... is
the described black-fruited mountain-ash" (Michurin 1948a: 142).
It is important to add that within the Baltic States and St.
Petersburg/Leningrad—the area essentially connected with Germany as far as
propagule exchange—large-fruited aronia was first encountered only at the
end of 1940's. We never succeeded finding any information about
black-fruited aronia in the German literature of the time.
I.V. Michurin recommended to use the new cultivated plant for
agricultural shelter plantings and then process the fruit crop for a variety
of technical applications as well as "dessert preserves in those
harsh-climate areas where there is lack of other fruit." From the start of
the 20th century, black-fruited aronia makes its way to the list of trees
and shrubs with edible fruit. For example, V.V. Pashkevich (1912) said
aronia fruit was suitable for jams. However, the brevity of that note makes
us think he might not know the plant very well; it is also remains unclear
whether he meant the wild species or new large-fruited aronia.
All our attempts to interview veterans of introduction business and
browsing through literature sources did not yield any information about
propagation and spread of Michurin's aronia early in the 20th century. Even
the staff of the Pomology Research Institute named after Michurin (in
Michurinsk) could not provide any such information at our request. It
appears that up until 1930's Michurin's aronia was not much promoted.
Documented history of its introduction starts only in 1935
I.V. Michurin died in 1935.
when Mikhail A. Lisavenko
1897-1967, a renown
Russian horticulturist, Director of Altai Horticultural Research Institute
in Gorno-Altaisk/Barnaul (1932-1967)
brought a few cuttings from Michurinsk
The capital of Altai Republic within the Russian
Federation, formerly known under names of Oirot-Tura and Ulala
cuttings successfully overwintered outside, under the snow cover and became
the source of extensive plantations of aronia in Siberia. In the 40's the
Altai Research Institute (then called Experimental Station) started sending
out seed and seedlings of Michurin's black-fruited aronia out to many
different destinations within this country. In 1940, 10 seedlings from the
Altai Experimental Station were planted in Vilgort, Komi Autonomous Republic.
The seedlings proved to be hardy there and started to fruit
after 4 years (Charochkin 1954).
In 1947 aronia from Gorno-Altaisk was introduced to Kaluga Oblast,
central European Russia, near Moscow (Myatkovsky, Telyukov 1966), and in
1958 a first commercial plantation was started there in Bukhlovka State
Farm. By 1969, there were 568 hectares (1,420 acres) of plantations around
Kaluga (Myatkovsky 1970).
Also in 1947 the Gorno-Altaisk Experimental Station forwarded a load of
aronia seedlings to Lesnoye [Forest] State Farm in Leningrad Oblast
(Vasilchenko, Protsenko 1967). This institution became a source of aronia
for the European Northwest of the USSR. Large shipments of seedlings were
sent from there to Latvia, Lithuania, and Belorussia, as well as Kalinin
(or Tver) Oblast and Vladimir Oblast (Shchukina 1967).
In 1948 cuttings of Michurin's aronia were forwarded from Leningrad to
A town near the City of Murmansk, beyond the Polar Circle, at
67° N, on the White Sea (or Beloye More) Coast
and first kept in a
greenhouse there. Two-year-old cuttings of the second greenhouse generation
were planted out near Apatity Railroad Station in 1954. They started
fruiting in 6 years. According to B.V. Kester (1970), aronia of the
Leningrad origin overwinters in Apatity with live terminal buds, flowers
yearly, though fruiting occurs only during favorable years, and only 20-30%
of fruit mature to ripeness.
With the material from Lesnoye, Leningrad, new plantations of aronia
were stated in Zelenogorsk (Leningrad Oblast) in 1954 (Ignatenko 1965). In
1958 aronia from Lesnoye was also introduced to Kharyus in Estonia, and by
1965 the plant was cultivated in four different state farms in Estonia (Kask
1971). The population in Petrozavodsk, Karelia, northwestern Russia also
originates from Lesnoye State Farm in Leningrad (Izergina 1969, 1971).
At the same time the area under the Siberian plantations continued to
grow. A new plantation in Bakchar Branch of Tomsk Agricultural Experimental
Station was started in 1957 (Gidzyuk 1966). Yet another one was developed in
Tavricheskiy [Taurida] Fruit Farm in Omsk Oblast in 1964 (Ryzhkov
The Gorno-Altaisk Horticultural Experimental Station remained the major
source of the introduction material. In 1958 seedlings from the Station were
introduced to Aurgaz and Gafura Districts in Bashkiria
Now Bashkortostan, a
republic within the Russian Federation situated in the southern Urals Mts.
and south of the Urals
(Barabash 1960; Suslov 1969). In 1959 the Altai
Station forwarded the first shipment of Michurin's aronia seedlings to
nurseries and teaching/experimental gardens of Udmurtia
A republic within
the Russian Federation in the western part of the Central Urals
became one of the most widespread cultivated fruiting shrubs here. According
to M.G. Kontsevoy (1974, 1975), as of 1970, there were three state farms in
Udmurtia with plantations of aronia covering 40 hectares (100 acres)
each; and the total area in Udmurtia was 221 hectares (552 acres)
. In 1959
fist seedlings of aronia were planted in Kirov (or Vyatka)
seedlings successfully took, became acclimated, and in four years the first
commercial plantation was already underway. By 1974, the area under aronia
in Kirov Oblast was 500 hectares (1,250 acres)
(Cheglakov and others,
It is not known when Michurin's aronia was introduced to Ukraine. There
is a brief note in Musich & Alexeyenko (1978) about aronia there before
World War II, though there is no exact dates or sources mentioned. Indeed it
is not even known whether that was Michurin's aronia or its wild ancestor,
which, as we have said here before, had been grown in Kharkov and Kremenets
as early as the beginning of the 19th century.
On the Island of Sakhalin (in the Russian Far East), in the State Farms
Yablochnyy and Pyatirechenskiy [Apple Orchard Farm and Five-River Farm],
there grows aronia originating from Altai and also from Leningrad material
(Voronova 1967, Shchukina 1967).
Taking all of this information into account, we may state with
confidence that all of cultivated aronia in this country originates from
Michurin's nursery. The area of cultivated aronia covers a significant part
of the USSR (Fig. 1), with a tendency for further expansion. For example, it
has been being introduced in Moldavia and Belorussia's (Bibikov, Kramnik
1974; Zhungietu 1975), in the Baltic States, and Northern Caucasus
(Myatkovsky 1970). Michurin's aronia is also successfully cultivated in
private gardens and teaching/experimental stations in Arkhangelsk Oblast
(about 64° N), coming there from Petrozavodsk and Vologda; in Volgograd
Obl. (starting from 1963; introduced from Kuibyshev (or Samara)); in
Astrakhan Obl. (about 46° N, at the mouth of the Volga River flowing
into the Caspian Sea) (introduced in 1970 directly from Michurinsk).
Fig. 1. Range of cultivated aronia in the USSR, according to literature
own observations, and oral communications.
1— areas where
aronia is considered a cultivated plant with acclimated cultivars
2—locations according to literature sources
according to herbarium records
4—generalized boundary of
II. Flowering of cultivated aronia
In order to reveal causes of low variability of the cultivated aronia,
we conducted experiments in June 1978, which included isolation, castration,
and artificial pollination of flowers. The experiments were conducted in the
Central Botanical Garden USSR Acad. Sci. using 5-year-old cultivated plants.
At the stage of flower buds, the inflorescences were isolated with 3-layer
mesh or oil-paper bags. There were 12 isolated inflorescences in each of the
following 7 lots in the experiment:
- Castrate flowers and remove stigmas
- Castrate flowers without removing stigmas
- Castrate flowers and pollinate with Chaenomeles japonica pollen
- Castrate flowers and pollinate with pollen taken from the same flower
- Castrate flowers and pollinate with pollen from the same inflorescence
- Castrate flowers and pollinate with pollen from the same specimen
- Castrate flowers and pollinate with pollen from the same population
Two different control lots were isolated intact flowers and
flowers subject to open cross-pollination.
Such experiments conducted earlier had arrived to controversial
results. For example, Bibikov & Kramnik (1974) did not obtain any fruit
upon pollinating castrated flowers with alien pollen; at the same time,
Kuzmina (1975) reported fruit formation upon pollination with alien pollen
and upon treating stigmas with various stimulators.
In 1978 aronia started to flower on May 29, finishing on June 21, with
a peak of flowering during June 4-10. There were on average 26 flowers per
inflorescence. Flowering of a single inflorescence continued 10-13 days (4-5
days for each flower).
Bibikov & Kramnik (1974) observed some specimens with double
flowers in aronia populations. In our plantings we did not have any
specimens whose all flowers were double. When found (in 3 cases total), some
double flowers were present within inflorescences with normal flowers.
Lot 1 (castrate flowers , remove stigmas) yielded only 6% fruit
Lot 2 (castrate flowers, intact stigmas) yielded 58%.
Lot 3 (castrate flowers, pollination with Chaenomeles japonica) yielded
Lots 4, 5, 6 yielded 95-99%.
Result for Lot 7 is not mentioned.
Control Lot 1 (isolated intact flowers) produced 68 % fruit.
Control Lot 2 (free pollination) yielded 79%.
These results demonstrate that aronia is capable of autonomous,
spontaneous apomixis; however, with participation of pollen (either own or
alien), fruit formation becomes more abundant. Taking into account the lack
of variability in the progeny, we have to conclude that there is no real
amphimixis in our aronia.
On the contrary, in A. melanocarpa within its natural range (North
Carolina), similar experiments (isolation, castration, and artificial
pollination) did not yield any evidence of apomixis (Hardin 1973). Moreover,
in the northern part of its natural area (Nova Scotia), A. melanocarpa was
found not to set any fruit even when self-pollinated, that is, behaving as a
species with cross-pollination (Hall et al.1978). At the same time,
A. arbutifolia is known to be apomictic (Hardin 1973); apomixis is also rather
common in Sorbus. Therefore, apomictic character of Michurin's aronia is not
something completely unexpected.
III. Chromosome number in cultivated aronia
Fig.2. Sets of chromosomes in metaphase plates of aronia.
collected in Moscow.
b. Seed collected in Barnaul.
All literature sources (Moffett 1931, Sax 1931, Hall et al. 1978)
report 2n=34 for A. melanocarpa as well as A. arbutifolia and
A. x prunifolia. Moffett (1931) detected 2n=68 in
A. arbutifolia cultivated in
We counted chromosomes in cultivated aronia originating from Moscow,
Riga, Barnaul, and Sakhalin.
The following method was employed. Seeds packed in tissue bags, 40 g
each, were soaked in water at room temperature for 24 hrs. Then the seeds in
bags were stratified in damp peat moss (to prevent desiccation) for 4 months
at 0 to +1°C. Stratified seeds were placed in Petri dishes on moist
filter paper at room temperature for germination. In order to accumulate
metaphase plates and obtain good chromosome spread in cells, roots 1-1.5 cm
long were placed in 0.5% colchicine solution at 3-5°C for 24 hrs. and
upon that fixated in Karnua solution for another 24 hrs. The material was
washed twice in 80% alcohol and then preserved in 60% alcohol until the
start of the count. Roots were stained in 5% acetocarmin, while the solution
was heated to boiling for 5-8 min. In order to remove excess stain, roots
were then kept for 10 min. in 45% acetic acid. Chromosome counts were made
on temporary slide mounts, photographed with Amplival Microscope equipped
with mf-matic device for microphotography.
All metaphase plates demonstrated 2n=68 (Fig. 2), which showed that
Michurin's aronia cultivated in the USSR is an apomictic tetraploid
IV. On probable mechanisms of cultivated aronia formation
It is a known fact that both polyploidy and apomixis can either be
induced by various factors or be of purely genetic nature. It is also known
that polyploidy often results in apomixis. Michurin practiced crossings
widely, including those of very remotely related species or those completed
with pollen mix originating from a few species. Alien pollen could act as a stimulant
triggering a transformation to polyploidy and apomixis. This scenario seems
quite probable—especially if we consider prevalence of polyploidy and
apomixis in the subfamily Pomoideae.
Yet polyploidy may result only in some (insignificant or even
negligible) changes in size of plant organs, whereas the difference between
the cultivated tetraploid aronia and its wild diploid ancestor is mostly
qualitative (Skvortsov, Maitulina 1982). Therefore, it would be reasonable
to assume that, in addition to changes in ploidy, some more important genome
changes took place during the formation of the cultivated aronia. This could
be, for example, a replacement of a chromosome(s) within a homologous
pair(s) as a result of interspecific crossings with subsequent recombination
during a hybrid split (let's remember that Michurin mentioned "selection
within three generations"). Translocations could also take place, either
with connection to hybridization or not. Any heterozygosity could have
become permanent due to apomyxis.
Examples from another genus, Oenothera (Onagraceae), demonstrate that
rather significant and permanent morpho-physiological changes in plants that
are based on structural genome changes can emerge quite rapidly. Introduced
from North America to Europe at the late 16th — early 17th century,
representatives of this genus gave rise to new species unknown in North
America, such as O. erythrocepala Borb. and O. rubicaulis Klebahn. It is
difficult to interpret the new species formation as a result of hybridization
with the European species, because these plants don't exhibit any characters
that appear to be inherited from a European parent or intermediate between
European and American plants.
Apparently, both in the case of Oenothera and Aronia,
the new species
formation has been triggered by drastic environmental changes, once plants
found themselves on a different continent. However, mechanisms of the
hereditary fixation of the changes have been different. There is no apomixis
in Oenothera; instead there is a particular genus-specific cytogenetic
mechanism that provides for the constancy of characters.
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Translation I. Kadis
30 December 2011