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Wild grapevine (Vitis sylvestris C.C.Gmel.) wines from the Southern Caucasus region

Authors:
  • Scientific Research Institute of Viticulture and Wine-making of Azerbaijan

Abstract

Grapevine domestication took place in the Caucasus area known as the Cradle of Viticulture, within or near the geographical area known as the Vavilov Triangle. The phytogenetic resources of Vitis sylvestris C.C.Gmel. have been previously collected and characterized, but the study on micro vinifications of wild grapevines from the Caucasus is new.In the present document, seven grape samples from female individuals of wild grapevine growing in the South Caucasus region were investigated to assess their oenological profile.Wine samples were obtained from the grapes collected from various populations of Armenia, Azerbaijan and Georgia in October 2013 and fermented by the native yeasts.Parameters determined in the wines were, among others, concentration of ethanol (3.63% – 10.15%), pH (3.30 – 4.20), total acidity (1.2 – 10.7 g/L of tartaric acid), total polyphenol index (1.81 – 89.8), and color intensity (2.59 – 20.76). This wide range values is due to the different environmental conditions, the level of ripeness of harvested grapes and their genetic diversity. These data were compared with those obtained in micro vinifications of wild grapevines in Western Europe and wines of several international cultivars.The results of our research demonstrated, that the must of wild grape could be used to improve traditional wines giving to them more coloration.
Wild grapevine (Vitis sylvestris C.C.Gmel.) wines
from the Southern Caucasus region
David Maghradze1,2, Gagik Melyan3, Vugar Salimov4, Ramazi Chipashvili5, Monserrat Íñiguez6, Pilar Puras6,
Elena Melendez6, Ramón Vaca7, Carlos Ocete8, Diego Rivera9, Concepción Obón12, Jose Manuel Valle10,
Alvaro Rodriguez-Miranda10, Osvaldo Failla11 and Rafael Ocete8
1 Scientific – Research Center of Agriculture, Marshal Gelovani Ave. 6, 0159 Tbilisi, Georgia
2 Faculty of Viticulture and Winemaking, Caucasus International University, Chargali Str. 73, 0141, Tbilisi, Georgia
3 Armenian Academy of Viticulture and Wine-making, Pushkin str. 33/10, 375002, Yerevan, Armenia
4 Institute of Viticulture and Winemaking, Mekhtiabad vil. Absheron distr. 0118, Baku, Azerbaijan
5 Institute of Horticulture, Viticulture and Oenology, Agricultural University of Georgia, 0159,
David Aghmashenebeli Alley 240, Tbilisi, Georgia
6 Estación Enológica de Haro, Av. Bretón de los Herreros, 4. 26200. Haro, La Rioja, Spain
7 Batle Winery, Palma de Mallorca, Camí de Coanegra, s/n, 07320 Santa Maria del Camí,
Illes Balears, Spain
8 University of Seville, San Fernando 4, 41004, Seville,Spain
9 Departamento de Biología Vegetal. Facultad de Biología. Universidad de Murcia,
Campus de Espinardo, 30100 Murcia, Spain.
10 Laboratorio de documentación geométrica del patrimonio. University of Basque Country (UPV/EHU),
Centro de Investigación Micaela Portilla, 01006 Vitoria – Gasteiz, Spain
11 University of Milan, Via Celoria 2, 20133 Milan, Italy
12Departamento Biología Aplicada, EPSO, Universidad Miguel Hernández de Elche,
30312 Orihuela (Alicante), Spain
*Corresponding author: drivera@um.es
Grapevine domestication took place in the Caucasus area known as the Cradle of Viticulture, within or near the
geographical area known as the Vavilov Triangle. The phytogenetic resources of Vitis sylvestris C.C.Gmel. have
been previously collected and characterized, but the study on micro vinifications of wild grapevines from the
Caucasus is new.
In the present document, seven grape samples from female individuals of wild grapevine growing in the South
Caucasus region were investigated to assess their oenological profile.
Wine samples were obtained from the grapes collected from various populations of Armenia, Azerbaijan and
Georgia in October 2013 and fermented by the native yeasts.
Parameters determined in the wines were, among others, the concentration of ethanol (3.63 % - 10.15 %), pH
(3.30 - 4.20), total acidity (1.2 - 10.7 g/L of tartaric acid), total polyphenol index (1.81 - 89.8) and colour intensity
(2.59 - 20.76). This wide range of values is due to the different environmental conditions, the level of ripeness of
harvested grapes and their genetic diversity. These data were compared with those obtained in micro vinifications of
wild grapevines in Western Europe and wines of several international cultivars.
The results of our research demonstrated, that the must of wild grape could be used to improve traditional wines
giving them more colouration.
micro vinification, river-bank forests, ethnobotany, Vitis vinifera L. ssp. sylvestris (Gmelin) Hegi, wine, Caucasus
A B S T R A C T
K E Y W O R D S
Received: 15 June 2020
y
Accepted: 15 October 2020
y
Published: 30 0ctober 2020
DOI:10.20870/oeno-one.2020.54.4.3720
VINE AND WINE
OPEN ACCESS JOURNAL
849
OENO One 2020, 54, 4, 849-862 © 2020 International Viticulture and Enology Society - IVES
INTRODUCTION
The Eurasian wild grapevine, Vitis sylvestris
C.C. Gmel., constitute the dioecious parental of
Vitis vinifera L. cultivars, which are usually
hermaphrodites (Rivera and Walker, 1989; This
et al., 2006; Zohary, 2000). The Eurasian wild
grapevine has received very diverse taxonomic
treatments, from the rank of variety to one of the
species. This implies the use of the subsequent
valid names, depending on the accepted level:
Vitis vinifera var. sylvestris Willd., V. vinifera
subsp. sylvestris (Willd.) Hegi or V. vinifera
subsp. syl vestr is (C.C. Gmel.) Hegi, an d
V. sylvestris C.C. Gmel. (Ferrer-Gallego, 2019).
Fossils of this autochthonous vine for Eurasia
appear within sediments dated from the end of
the Pliocene (Sémah and Renault-Miskovsky,
2004). At present, these wild populations are
disseminated in natural ecosystems from the
Iberian Peninsula to Hindu Kush (Arnold et al.,
2002). Some populations of this liana can be also
found in the African Maghreb (Ocete et al.,
2007). Their main habitats are river-bank forests,
river mouths, flood plains, colluvial positions on
the slopes of hills and mountains and coasts
between the para llels 4 N (Rhine River,
Germany) and 30º N (Ourika River, Morocco)
(Iriarte et al., 2013). In such places, soils are
often renewed by flooding (Arnol d, 2002;
Maghradze et al., 2010).
Pallas (1799 - 1801), a German naturalist at the
service of Empress Catherine II of Russia,
reported the presenc e of countless wild
grapevine populations in the Southern Caucasus.
There were several individuals with large logs,
some of them with the thickness of a ship’s mast;
their branches climbed on the surrounding trees.
Bunches of grapes were harvested by the
inhabitants of the region, sometimes, when the
entire grape became raisin after winter frost, in
the spring season. Eyriés (1841) indicated that
the grapevine grows in the gullies and plains of
Southern Caucasus as in their primitive
homeland. Thus, suggesting this area to be part
of a centre of domestication for grapevine, which
is consistent with recent molecular data: “The
close association of Georgian wild grapevines
with Georgian cultivated accessions strongly
supports their inv olvement in the initia l
domestication of grapevine” (Riaz et al., 2018).
The Caucasus became even more relevant for
understanding Vitis sylvestris diversity after the
choice of a neotype for this taxon by Ferrer-
Gallego et al. (2019) who designated the
specimen collected in Georgia (Alazani river
basin, Jumaskure, 41°21.588′ N, 46°35.934′ E)
by Ia Pipia, which is preserved in the Herbarium
of the Institute of Botany, Ilia State University
(TBI barcode TBI1052417!).
The South Caucasus region is situated between
the Black and Caspian seas, across several
countries, notably Armenia, Azerbaijan and
Georgia, and is an important refuge area for
numerous plant species including sweet chestnut,
walnut and wild grapevine (Aradhya et al., 2017;
Krebs, 2019; Ramishvili, 1988; Ramishvili,
2001). Several wild relatives of domesticated
fruit species are present there in relic habitats in
the Greater Caucasus mountain range (Huglin
and Schneider, 1986; Vavilov, 1931). It
constitutes the territory with the highest Eurasian
grapevine diversity (wi ld and cultiv ated)
(Haxthausen, 1856; Kolenati, 1846; Negrul,
1938; Vav ilov, 1926) and it is part of the
grapevine’s “Fertile Triangle” or “Vavilov´s
Triangle” (Figure 1) (Robinson et al., 2013). The
South Caucasus region has been postulated as
the cradle of viticulture and winemaki ng
(McGovern, 2003; 2004, McGovern et al., 2017;
Zohary, 2000).
In South Caucasus Region wild grapevine climbs
on numerous tree and shrub species in open
woodland ( Ocete et al., 2018). Uses of
Caucasian wild grapevine include medicine;
agriculture (pollination of female cultivars) and
food (male flowers flavour ing wines i n
Azerbaijan, and unripe fruits (verjuice) in
marinades and special sauces (Maghradze et al.,
2015b).
The Eurasian wild grapevine is considered a
threatened plant genetic resource due to the
overexploitation of riverine forests, and the
establishment of orchards and public works. The
importation of fungal diseases from North
America, such as downy and powdery mildews
strongly reduced natural populations (Ocete et
al., 2015). Furthermore, after Phyll oxera
infestation, there was a massive incorporation of
North American Vitis species in Eura sian
vineyards. They were used as root-stocks and in
genetic improvement projects addressed at
obtaining direct producer hybrids (French-
American hybrids). Both kinds of plants showed
a heavy invasive character as feral plants in wild
habitats, highly competitive in the same habitats
David Maghradze et al.
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850
where lived autocht honous Eurasian wild
grapevine (Ocete et al., 2007; Terpó, 1974).
Wild grapevine reproduces mainly by seed,
differi ng from the established vegetative
propagation of cultivars (Iriarte et al., 2013;
Revilla et al., 2010), and presents a higher level
of genetic diversity, particularly in South
Caucasian Region (Pipia et al., 2015). Genetic
studies including haplotype distribution based on
plastid DNA sequences show high levels of
variation in wild grapevine (V. vinifera subsp.
sylvestris) from the Greater Caucasus region
(Pipia et al., 2012). In natural wild populations
mutations affecting male vines can originate
hermaphrodite individuals (Picq et al., 2014).
Early farmers selected hermaphrodite
grapevines, presumably due to their higher
production of grapes and easier cultivation, to
establish the first vineyards outside river-bank
forests (Forni, 2006, 2012; Scienza, 2004; This
et al., 2006). However, some degree of dioecy
coexisted in cultivation. The South Caucasus
Region houses numerous female cultivars (97
out of 725 for the whole area, 53/414 in Georgia,
22/144 in Azerbaijan, and 22/171 in Armenia)
(Negrul, 1970). In the years of intensive
development of viticulture in Azerbaijan, it was
carried out artificial pollination of functionally
female grapevine varieties (Ag shany, Khatuni,
Tavkveri, Nimrang and others) with pollen of
male inflorescences of wild grapes to enhance
the productivity of vineyards (Efendiyev, 1972).
© 2020 International Viticulture and Enology Society - IVESOENO One 2020, 54, 4, 849-862 851
FIGURE 1. The “Vavilov’s Triangle” and sampled localities.
Shulaveri-Shomu culture existed on the territory
of p resent-day Georgia, Azerbaija n and
Armenia. The culture is dated to the 6th or early
5th millennia BC and is thought to be one of the
earliest known Neolithic cultures. Some of the
first wine production artefacts were found in the
archaeological sites of Shulaveri Gora and
Gadachrili Gora in South Georgia with other
evidence of agricultural activities dated c.
8000 BP (Chilashvili, 2004; McGovern, 2003;
McGovern et al., 2017) (Figure 2). Archaeolo-
gical excavations in the Areni-1 cave complex in
south-eastern Armenia revealed installations and
artefacts dating to around 6000 BP that are
strongly indicative of wine production (Barnard
et al., 2011).
It is necessary to remark that liquid products
other than wine were obtained from grapes
during the Prehistory and Antiquity. Grape must
was used to improve ceramic pastes, at least,
from the Bronze Age and grape vinegar was a
very important food preserver used in beverages
such as the poscaconsumed by Roman
legions (Ocete et al., 2011c). The population of
ancient Azerbaijan used wild grapes in food.
Over time, local residents began to move wild
grapevine closer to its homes and cultivate it.
Remains of wild grapevine were found among
the rocks of the ancient Gobustan and in the
Khachmass region of Azerbaijan (Babayev,
1988).
The grapevine cross, or Saint Nino’s cross, is a
major symbol of the Christian Georgian
Orthodox Church. Saint Nino of Cappadocia,
who preached in Georgia in the 4th century AD,
is represented as a girl holding up a cross made
with shoots of grapevine tied using her own hair
(Maghradze et al., 2015a).
The Eurasian wild grape produces a rather
astringent, small fruit with numerous seeds,
hardly the kind of grape for making good wine.
Its sugar content is relatively low and acids are
high, as compar ed wi th th e domesticated
Eurasian cultivars, and the skin of its fruit is
tough (McGovern, 2003). Therefore, it could be
expected that wine obtained from these grapes
would differ in certain analytic parameters from
common wines.
An ampelography of selected native grape
varieties of the six cou ntries Azerbaijan,
Armenia, Georgia, Moldova, Russia and Ukraine
has been published. T he ident ification,
collection, characterization and conservation of
the diversity of grapevine genetic resources was
done 2004 - 2008 (Maghradze et al., 2012).
David Maghradze et al.
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FIGURE 2. Archaeological grape vine evidence.
A, Grape pips from ShulaveriGora (Georgia) c. 6000 BC (Tbilisi Archaeological Museum);
B, Large vessel with decorations imitating clusters of grapes supposedly used to have contained wine, c.
6000 BC (Tbilisi Archaeological Museum); Images: R. Ocete.
According to the philosophy of the COST FA
1003 Action: East-West Collaboration for
Grapevine Diversity and Exp loration and
Mobilization of Adaptive Traits for Breeding”
(2010 - 2014) an expedition to collect and
conserve plant genetic resources of grapevines
from the South Caucasian Region was carried
out in 2013.
Georgian cultivated and wild grapevine has been
described (Chkhartishvili and Maghradze, 2012;
Ocete et al., 2012) and genetically characterized
(De Lorenzis et al., 2015; Ekhvaia et al., 2014;
Imazio et al., 2013;Imazio et al., 2013), but not
the winemaking with wild grapevine of this
country, likely in Azerbaijan (Salimov and
Musayev, 2012) and Armenia (Melyan and
Gasparyan, 2012).
We believe it is important to make it clear that
wild grapevines in the Caucasus are a n
important genetic resource for all the reasons
above stated. The wil d grapevines of the
Caucasus have been studied and characterized
genetically and morphologically but there is a
lack of data of the characteristics of the wine
they provide.
The wild grapes have been vinified since ancient
times and are still used for this purpose both in
© 2020 International Viticulture and Enology Society - IVESOENO One 2020, 54, 4, 849-862 853
FIGURE 3. Harvest of wild grapes and habitats.
A, Harvest of wild grapes in Guruchai River (Azerbaijan); B, Ripe wild grapes, Guruchai River; C, Ripe
wild grape from Ktsia River (Georgia); D, Fruiting wild grapevine in Guruchai River (Azerbaijan).
E, Harvesting climbing grapevine (Georgia). F, Climbing wild grapevine and supporter (Azerbaijan).
Images: D. Maghradze and V. Salimov.
the study area and in other places where wild
grapevine grows (for example in Sardinia).
For al l this, the ai m of this work is: to
characterize the wine that is obtained from wild
grapes harvested in the several populations of
Azerbaijan, Georgia and Armenia; to establish a
preliminary characterization on the oenological
potential of wild grapes within this geographical
area; to know better the likely compositions of
the wines produced bef ore grapevine
domestication.
MATERIALS AND METHODS
1. |Sampling
Harvest of grapes took place at the second
middle of October 2013 in Armenia, Azerbaijan
and Georgia in the wild grapevine populations
free of the presence of feral cultivars and
American root-stocks (Figure 3).
The coordinates of the different populations
sampled along river-bank forests and flood
plains are shown in Table 1 and Figure . These
lianas climb on several sp ecies o f the
accompanying vegetation, such as Carpinus
betul us, Co rnus m as, Co rylus av ell ana,
Cr ataegus caucasica, Mesp il us germanica,
Paliuru s spina- christi , Prunus divaric ata,
Pu ni ca gr an at um , C yd on ia ob lo ng a, Py ru s
ca ucasica, Quercus iberica, Salix capreae,
Ulmus minor among others (Ocete et al., 2018).
All-female plants had red suborbicular berries,
with diameter inferior to 1 cm. The skin of the
grape is blue-black or dark red-violet (Table 1,
Figure 3). The surface is covered with a thick
wax layer.
2. Wine production and analysis
Bunches containing a considerable proportion of
ripe grapes were selected among those available
for harvest. High heterogeneity in the fruit set
and ripening process observed in the same
cluster (millerandage) is characteristic of wild
grapevine populations (Trad et al., 2017). The
removal and separation of ripe grape berries
from t he s tems (des temming) were done
manually. Of each sample, 50 berries preselected
as ripe were weighed t o calculate w hat
percentage is transformed into must. Only ripe
berries were pressed using manual machinery.
Given the small number of grapes available, only
one sample from each locality (Table 1) was
fermented, no replicas were made. The fermen-
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854
* We follow the rules of IPGRI-UPOV-OIV (1997). ** Given that wild grapevine plants are climbing on different tree supports, the upper branches receive abundant light (Ocete et al.,
2018). ***Climate data from NOAA (2020): Guba for Guruchai River, Bolnisi for Ktsia River, Gardabani for Mtkvari River and Odzun for Debet River. Data on temperatures are not
available.
TABLE 1. Geographic information of wild grape populations from their natural habitats in South Caucasus countries and characteristics of their grapes.
Place/ Population Guruchai River 1 Guruchai River 2 Guruchai River 3 Guruchai River 4
Latitude 41º24'01" 41º26'03" 41º28'09" 41º27'43"
Longitude 48º26'37" 48º33'41" 48º33'59" 48º35'25"
Berry skin colour* Blue-black Dark red-violet Blue-black Blue-black
Berry shape* Round Round Round Round
Habitat
Remains of Quercus iberica forest
in anthropized habitats
Populus alba riparian forest
Remains of Quercus iberica forest
in anthropized habitats
Remains of Quercus iberica forest
in anthropized habitats
Slope orientation, altitude and sun exposition** Slightly north facing, 681 m a.s.l, sheltered Slightly north facing, 407 m a.s.l, sheltered Almost flat, 384 m a.s.l, sun-exposed Almost flat, 346 m a.s.l, sun-exposed
Average precipitation (mm)*** 416 416 416 416
Wine 5 Wine 6 Wine 7
Armenia
Place/ Population Ktsia River 1 Mtkvari River 1 Debet River 1
Latitude 41º29'22" 41º22' 43" 41º07'16"
Longitude 44º40'51" 45º03' 25" 44º45'16"
Berry skin colour* Dark red-violet Dark red-violet Dark red-violet
Berry shape* Round Round Round
Habitat
Punica granatum
and Crataegus riparian thicket
Punica granatum
and Elaeagnus riparian thicket
Quercus iberica fores
with Fraxinus and Acer
Slope orientation, altitude and sun exposition** North facing, 421 m a. s.l., shaded West facing, 277 m a. s. l., sun-exposed Steep slope west facing, 652 m a. s. l., sun-exposed
Average precipitation (mm)*** 495 370 650
Azerbaijan
tation was carried out in the laboratory in glass
jars, the first four weeks, and then transferred to
bottles for transport, with the own yeasts that
carried the berries (spontaneous fermentation),
for a maximum of 15 da ys, with a fix ed
temperature of 20 ºC and daily stirring of the
must with the skins of the berries. There was no
addition of potassium metabisulfite. The samples
were analyzed following the methods proposed
by the OIV (2015) in a laboratory accredited
under Quality Standard 17025 (ISO 2019), as
follows:
- Ethanol: Near Infrared (NIR) (SpectraAlyzer
WINE, ZEUTEC).
- pH and total acidity: Automatic potentiometry
(Win elab Analyzer, FOODLAB-CD R,
Florence, Italy - Tecnología Difusión Ibérica,
Barcelona, Spain).
- Tartaric acid: Enzymatic (Cetlab 600,
Microdom, Taverny, France - Tecnoloa
Difusión Ibérica, Barcelona, Spain).
- Total polyphenol index: UV spectrometry
(LAMBDA 265 PDA UV/Visible Spectropho-
tometer, cuvettes (1 mm pathlength), Perkin
Elmer, Waltham, Massachusetts, USA).
- Colour intensi ty: UV-VIS spectr ometry
(LAMBDA 265 PDA UV/Visible Spectropho-
tometer, cuvettes (1 cm pathlength), Perkin
Elmer, Waltham, Massachusetts, USA).
- L- Malic acid and volatile acidity: Enzymatic
(Cetlab 600, Microdom, Taverny, France -
Tecnología Difusión Ibérica, Barcelona, Spain).
- Reducing sugars: Autoanalyzer FCSA Q05
with Quaatro 39 (SEA L, Nord erstedt,
Germany - AXFLOW, Arsta, Sweden).
3. Comparison
To determine relationships within the wines
obtained we calculated the pairwise differences
between samples in form of a dissimilarity
matrix.
The crude matrix consisted of 8 variables
(ethanol content (% vol), total acidity (g/l), pH,
tartaric acid (g/l), L-malic acid (g/l), colour
intensity, total polyphenol index and reducing
sugars (g/l)) and 18 units (defined using mean-
sd, mean, and mean+sd values for each of the
6 samples). The matrix of chemical parameters
was used to compute a dissimilarity matrix using
DARwin V.6.0.17 (2018-04-25) (Perrier et al.,
2003; Perrier and Jacquemoud-Collet, 2006).
The c hi-square dissimilarity index wa s
calculated. This measure expresses a value xik as
its contribution to the sum xion all variables and
is a comparison of unit profiles [1].
where dij: dissimilarity between units iand j;
xik, xjk: values of variable kfor units i and j;
xi., xj .: mean for units iand j; x.k: mean for
variable k; x..: overall mean. K: number of
variables.
To realistically represent individual relations, a
hierarchical tree was constructed to describe the
relationships between units (samples) based on
the common agglomerative he uristic th at
proceeds by successive ascending agglome-
rations. For updating dissimilarity during the
tree construction, the Ward criterio n was
adopted, which searches at each step for a local
optimum to minimize the within-group or
equivalently to maximize the between-group
inertia. For the graphic representation, we have
opted for the software Figtree version 1.4.3.
(Rambaut, 2014). Analytical data of comparison
samples were obtained from De Gianni (2015)
(Nero d’Avola wine), Fogaça and Daudt (2012)
(Brazilian V. vinifera cultivars), Revilla et al.
(2016) (Spanish V. vinifera cultivars), Ocete et
al. (2011b) (Spanish wild grapevine wines),
Kang et al. (2008) (Traditional Korean wines are
made by adding rice to grape juice and adding
yeast), V. rotundifolia cultivars (Morris and
Brady, 2004; Talcott, 2004).
RESULTS AND DISCUSSION
The must yield per kilogram of grapes harvested
was situated between 16-17 %, due to the low
proportion of pulp in the fruits. A wine volume
of less than 250 ml was obtained in each of the
micro fer mentations, so t he me thod of
distillation with electronic densitometry was not
applicable to calculate the ethanol concentration
(v/v). Overall, the ethanol content measurement
results were extremely low for a beverage that
could be called wine (Table 2). This may be due
to a low sugar content in the grapes.
Given that between the wine production in
Georgia and the analysis in Spain, a period of
several weeks elapsed (c. 40 days), it is likely
that spontaneous ma lolactic fermentatio n
occurred, which would explain why tartaric and
2
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© 2020 International Viticulture and Enology Society - IVESOENO One 2020, 54, 4, 849-862 855
malic acids represent only up to 50 % of total
acidity.
The fact that the grapes have been fermented
with local natural yeasts can influence the
analytical characteristics of the experimental
wines. Therefore, the differences between the
wines are due not only to different origins and
environmental conditions but to different yeasts
as well.
Data on micro vinifications (Table 2) can be
summarized as follows:
1. Azerbaijan
Wine 1 (Guruchai River 1). After fermentation,
the percentage of ethanol recorded in this sample
was 5.78%. This wine had good total acidity and
showed a normal concentration of tartaric acid
(Almela et al., 1996). The colour intensity was
very low, similar to a rosé wine.
Wine 2 (Guruchai River 2). This wine showed a
higher percentage of ethanol, 10.15 %. It is the
maximum found in this region of the South
Caucasus. Total acidity is adequate. The total
polyphenol index is high, the colour intensity is
good, 10.60 (it could be appropriate for a good
quality red wine obtained from cultivars).
Wine 3 (Guruchai River 3). This wine showed a
lower concentration of ethanol, 4.62 %. It has a
low concentration of tartaric acid. The total
polyphenol index could not be carried out due to
the small volume of the sample.
Wine 4 (Guruchai River 4). This sample has a
high total acidity, a low to normal amount of
tartaric a cid an d only 5.04 % ethan ol
concentration. The intensity of the colour and the
polyphenol index are normal according to its
maturity level.
2. Georgia
Wine 5 (Ktsia River 1). The concentration of
ethanol is 5.21 % vol. The intensity of colour
and the polyphenol index present very good
values. In this case, the phenolic maturity has
been more in advance than the technological one
as suggested by the sugars/acidity values ratio.
Wi ne 6 (Mt kvari River 1). The ethanol
concentration is 7.2 % vol. The colour intensity
and polyph enol index present de cidedly
acceptable values.
3. Armenia
Wine 7 (Debet River 1). The berries of this
sample were so small, and with hardly any pulp,
that barely any must volume was achieved and
several determinations could not be completed. It
showed the lowest percentage of alcohol of all
microvinifications. Due to the few parameters
determined (Table 2), it is not included in the
comparison.
Considering all the results, analytical parameters
mainly fall within the range of variation of
cultivated grapevine wines. Ripeness level and
sugar content are highly influenced by the degree
of shade produced by botanical supporters (trees
and shrubs) and the rest of the accompanying
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856
TABLE 2. Wild grapes from South Caucasus countries: characteristics of their wines.
Notes: X: Average. σx: standard deviation. Inap., Inappreciable.
* For comparison with colour intensity of Vitis vinifera wines: cv Mencia (5.72 - 12.98 by Sudraud method and 16.43 - 17.21 by
Glories method) and cv Alicante Bouschet (12.16 - 24.43 by Sudraud method and 13.73 - 28.08 by Glories method) from AOC
Valdeorras, Galicia, NW Spain (Revilla et al., 2016); cv Merlot (between 4.3 - 11.0 by Glories method) from the Campahna
Gáucha and Serra Gáucha regions of Brazil (Fogaça and Daudt, 2012). In bold samples of group A Figure 4.
Wine 1 Wine 2 Wine 3 Wine 4 Wine 5 Wine 6 Wine 7
Values (X ± !x) Values (X ± !x) Values (X ± !x) Values (X ± !x) Values (X ± !x) Values (X ± !x) Values (X ± !x)
Ethanol (%) 5.78 ± inap. 10.15 ± inap. 4.62 ± inap. 5.04 ± inap. 5.21 ± inap. 7.2 ± inap. 3.63 ± inap.
pH 3.58 ± 0.05 3.31 ± 0.05 5.64 ± 0.05 3.50 ± 0.05 3.30 ± 0.05 4.20 ± 0.05 -
Total acidity (g/L tartaric acid) 5.3 ± 0.4 7.7 ± 0.4 1.2 ± 0.4 10.7 ± 0.4 8.2 ± 0.4 6.1 ± 0.4 -
L-malic acid (g/L) <0.10 <0.10 1.11 ± 0.22 <0.10 ± 0.22 0.90 ± 0.22 1.71 ± 0.22 <0.10
Tartaric acid (g/L) 2.30 ± 0.35 2.78 ± 0.35 0.59 ± 0.35 1.79 ± 0.35 3.24 ± 0.35 1.57 ± 0.35
Reducing sugars (g/L) 1 ± 0.5 1.5 ± 0.5 1.7 ± 0.5 4.9 ± 0.5 1.3 ± 0.5 1.8 ± 0.5 -
Total polyphenol index 18.1 ± 0.9 51.8 ±1.7 - 29.9 ± 0.9 56.50 ± 0.9 89.8 ± 0.9 -
Colour intensity* 2.59 ± 0.058 10.60 ± 0.058 4.85 ± 0.058 3.76 ± 0.058 20.19 ± 0.058 20.76 ± 0.058 -
Parameters
vegetation in natural habitats, such as river-bank
forests and flood plains (Ocete et al., 2018)
(Figure 3). The concentration of anthocyanins of
the ski n of the berries that will form the
pigmented polymers of red wines is also affected
by shade and weather (Esteban et al., 2001;
Fulcrand et al., 2006) and varies even in the
same cultivar along different harvests (Revilla et
al., 2009) and in wild grapevines (Benito, 2015;
Revilla et al., 2010; Cantos et al., 2017).
The ethanol percentage of normal samples varies
between 4.62 % and 10.15 % (the abnormal
sample 7 presented 3.63 %). The colour intensity
varies between 2.59 and 20.76. It is necessary to
remark that a wine is considered red, after the
malolactic fermentation, when its intensity of the
colour is 3.5 at least, for ins tance by the
Regulatory Council of the Denomination of
Origin Rioja (Spain) (Riojawine, 2019).
In general, ethanol levels and, sometimes, colour
intensity values in Caucasus wines from wild
grapevines are lower than those registered in
micro vinification with wild grape samples from
Sardinia (Ital y) (Lovicu et a l., 2009)and
Andalusia, La Rioja, Castille and León and
Navarre (Spain) (Ocete et al., 2011a; Ocete et
al., 2011b). In the case of Spain, the maximum
ethanol content was 14 %, registered in a sample
harvested in Cáceres province (Extremadura)
(Ayala et al., 2011) and the top colour intensity
was 26.4 determined on a micro vinification
from the Ega River lava province, Basque
Country) (Meléndez et al., 2015).
Concerning the classification, colour intensity
and total polyphenol index determine three main
groups (Figure 4) (cf. Table 2).
Group I is characterized by the highest values of
total polyphenol index, 50 - 90 (mean 66) and
colour intensity, 10 - 21 (mean 17.2). Total
polyphenols and colour intensity are lower and
similar for Groups II and II (17-31 for
polyphenol index and 3-5 for colour intensity).
Group II presents a lower tartaric acid content
(0.2 - 0.9 g/L) in comparison with Group I (1.2 -
3.5 g/L) and Group III (1.4 - 2.7 g/L). Group II,
also, presents an extremely low total acidity
(0.8 - 1.6 g/L) and a higher pH (5.6 - 5.7).
Finally, ethanol content was found not useful to
recognize groups. Group I (Figure 4) include
South Caucasian wild grapevine samples: two
from Georgia and one from Azerbaijan. Whose
compositions show similarities with some of the
wild grapevine samples from Spain (Ayala et al.,
2011; Ocete et al., 2011a; Ocete et al., 2011b),
Korean wines (Kang et al., 2008) and Vitis
rotundifolia wines (Morris and Brady, 2004;
Talcott, 2004).
Guruchai River samples 1, 3, 4, which form
clusters II and III, produced wines that have
shown similarities with those of Vitis vinifera
cultivars and most wild Eurasian grapevine
samples from Spain.
It is worth to highlight that, from a molecular
marker perspective, South Caucasian popu-
lations belong to chlorotypes C and D, whereas
© 2020 International Viticulture and Enology Society - IVESOENO One 2020, 54, 4, 849-862 857
FIGURE 4. Relationships among Caucasus wine samples.
Note: Ward’s minimum variance tree. A, B, C variants within each sample that were defined using mean-sd (A), mean (B), and
mean+sd (C) values for each parameter).
Spanish ones belong to chlorotype A (Arroyo-
García et al., 2006; De Andrés, et al., 2012).
All samples present r educing sugars not
transformed in et hanol, at different
concentrations. The high total polyphenol index
and high acidity could be assumed responsible
for the disruption of the normal action of yeasts.
However, these are not significantly different
from the levels in wines from cultivars .
Moreover, Wine 3 has 1.7 g/L of reducing sugars
and low polyphenols and acids content (Table 2).
Therefore, we cannot associate this level of
sugars with problems in fermentation due to the
total polyphenol index and high acidity.
At the time of carrying out the analyses, the
laboratory did not have the instruments for the
study of aroma, so these data are not available,
despite their interest. It would also be interesting
to produce more wine to perform a sensory
analysis. However, several points make it very
difficult: the extremely low number of grapes
produced by wild strains in their natural habitats
of South Caucasus during episodes of drought,
the inter-annual irregularity in the harvest and
the difficult access to some of the populations.
The use of wild grapevine has been frequented
for producing wine throughout history.
Currently, the Eurasian wild grapevine is in the
list of Endangered Plant Species of Georgia
since 1982 (Chemonics, 2000). In Azerbaijan,
people have always produced red and white
wines. An interesting wine is the so-called “gora
shara b”, traditi onal of the region Guba-
Khachmaz, Shaki-Zaqatala, Garabagh. For
making this wine people use cultivated and wild
grapes gathered in forest and riversides (Salimov
and Musayev 2012).
In the Azerbaijan Resear ch Institute of
Viticulture and Wine-making buds and pollen of
wild grapes are used as a flavour for preparing
flavoured dessert wine like «nectar». This wine
is characterized by a particular taste and unique
flavour (Amanov, 2001; Amanov, 2005).
From unripe berries of wild grapevine, local
habitants prepare healing juice, called «gora
suyu» or «gara suyu». This juice is successfully
used in the treatment of diabetes. Grapes contain
numerous polyphenols, including the stilbene
resveratrol, the flavanol quercetin, catechins, and
anthocyanins that have shown potential for
reducing hyperglycaemia, improving β-cell
function, and protecting against β-cell loss.
Therefore, with a low mean glycaemic index and
glycaemic load, grapes or grape products may
provide health benefits to type 2 diabetics
(Rasines-Perea and Teissedre, 2017; Zunino,
2009).
An infusion of fresh leaves of the wild vine is
widely used for the treatment of rheumatism (as
a bath), as well as for improving eyesight
(Damirov and Shukurov, 1985).
In Sardinia, a traditional wine is known as «vinu
de marxani» or vino de volpe” is made with the
wild grape. Until the middle of the 20th century,
shepherds of the mountainous area of Sulcis
made their own wine with these wild grapevines,
which they called vino de caoprai (Lovicu et al.,
2009; Lovicu, 2013). Therefore, it has been
traditional to make wine completely with wild
grapes.
The potential contribution of wild grapes
(wine 1, wine 2, wine 5) to lower the pH of the
must by increasing the acid content, facilitating
good wine conservation, is extremely limited by
the considerab le drop in alcohol that this
addition can produce. Red wild grape wines
(wine 2, wine 5 and wine 6), despite their high
polyphenolic content that could help improve the
preservation of base wines and add a higher
concentration of anthocyanins, are of little use as
improvers of wines made with cultivated
varieties, for the same reason.
CONCLUSIONS
The wild grapevine populations cited in the
present paper could be useful to make deeper
oenological studies, such as the analysis of
anthocyanin fingerprints. Wild grapevines with
fruits rich in colour could be used to intensify the
colour in red wines, as long as their low ethanol
content can be resolved.
These wines (wines 2, 5 and 6) for their content
in polyphenols could be used for improving the
conservation of organic wines.
It is desirable that in these countries the traditional
wine of the wild grapevine continues to be made
and eventually added to the conventional local
wines, which would confer certain original
characteristics to the wines from the domesticated
cultivars of the Eurasian grapevine.
Acknowledgements: This work was carried out
under the project COST FA1003 Action “East -
West Collaboration for Grapevine Diversity and
David Maghradze et al.
© 2020 International Viticulture and Enology Society - IVES OENO One 2020, 54, 4, 849-862
858
Exploration and Mobilization of Adaptive Traits
for Breeding” (2014-2018). We are indebted to
the persons helping during organization and
realization of the expeditions: Ekateri ne
Abashidze (Institute of Horticulture, Viticulture
and Oenology, Georgia), Akaki Modebadze and
Kakha Karala shvili (Lagodek hi Rese rve,
Georgia), David Maghradze and Ghuto Kiknadze
(Gardabani Reserve, Georgia), Shikh said
Akhmedov (Quba, Azerbaijan), Manvel Sukoyan
(Dilijan, Armenia).
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... and a high level of total polyphenols (about 80 g/L), suggesting the suitability of must from wild grape for prolonged winemaking process. More recently, a study of seven accessions of wild wines made with grapes harvested in the forests of the South Caucasus countries, including Armenia, Azerbaijan, and Georgia, also demonstrated the diversity of enological parameters, providing the idea that the must of wild grape could be used to improve traditional wines, resulting in color intensity [21]. This data also suggests that the possible combination of wild and domesticated grapes can make a wine suitable for long ageing. ...
... However, the wines made with wild grapes collected in the forests can only provide general information about the wild wines [21,22]. Various limiting factors, such as notuniform berry maturation, geographical differences in plant locations, and birds picking ripe berries, can limit the effective demonstration of the maximum enological potential of wild grapes. ...
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... The maintenance of the trait in V. vinifera through the domestication until today was probably not intentional because there is no literature reporting powdery mildew disease in Europe and Asia before the 19 th century. However, literature suggested that both natural and intentional grapevines selection took place in the last two centuries in the region (Ocete et al., 2012;Maghradze et al., 2020), when the pressure of E. necator on grape cultivation became evident due to pathogen introduction from North America. A recent selection could explain the high frequency of resistance haplotype 149-208 within the Caucasian cultivars. ...
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Vitis vinifera L. is the most cultivated grapevine species worldwide. Erysiphe necator,the causal agent of grape powdery mildew, is one of the main pathogens affecting theviticulture. V. vinifera has usually little genetic resistance against E. necator and grapeproduction is highly dependent from agrochemicals. The main purpose of this workwas the study and the mapping of the resistance to E. necator in the Caucasian V.vinifera germplasm. The biparental mapping approach was chosen to investigate thegenetic basis of the trait and two F1 populations were developed by crossing Shavtsitskaand Tskhvedianis Tetra with two susceptible V. vinifera varieties, Chardonnay andGlera. The phenotypic resistance of parental plants and offsprings was studied by usingleaf discs bioassays and evaluating the infection features during the pathogen life cycle.Caucasian cross parents showed a resistance to E. necator and the trait segregated intheir populations: the resistant genotypes delayed and limited the development ofpathogen mycelium, sporulation and conidia but they did not halt completely theinfections. A total of 184 seedlings of Shavtsitska cross population were genotypedthrough the Genotyping by Sequencing (GBS) technology allowing the development oftwo high-density linkage maps for the cross parents. QTL analysis revealed a majorresistance locus on Shavtsitska linkage group 13. Such a QTL was associated with areduced pathogen development as well as an enhanced plant necrotic response. TheQTL explained up to 80.15% of the observed variability and was restricted in aninterval of approx. 2.2 cM. The comparison with grape reference genome PN40024located the QTL at about 18 Mb from the top of the DNA sequence on chromosome 13and recombinants analysis restricted the locus in a region of 1.4 Mb. Some SSR locatedin the genomic region were used for genotyping the cross populations of the study and103 further Caucasian varieties. Resistance associated SSR alleles were shared amongShavtsitska, Tskhvedianis tetra, resistant seedlings and 22 Caucasian varietiessuggesting a widespread presence of the resistance trait in the Caucasian germplasm.The QTL isolated in Shavtsitska located in the region where the Ren1 resistance genecarried by several Central Asia V. vinifera was previously mapped. Further molecularanalysis is needed to confirm whether different genes in such region, that in thereference genome PN40024 is rich of resistance motifs, control the resistance to E.necator in grape accessions of different geographic origin. Meanwhile, our findings would provide new V. vinifera genetic sources for grape breeding programs aiming toobtain resistant elite cultivars.
... The female cultivar 'Angur Kalan' (syn. 'Nimrang') was found to be a major founder of local Armenian genetic resources (Table 3), which agrees with its common use as a female genitor in breeding programmes aimed at enhancing the productivity of vineyards during the development of viticulture in the South Caucasus (Maghradze et al., 2020). For practical reasons, renowned breeders like Bruno Bruni, Giovanni Dalmasso, Pierre Landot or Alberto Pirovano also recurrently used female cultivars as genitors in their breeding programmes. ...
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The South Caucasus is recognised as the primary Vitis vinifera L. (grapevine) domestication centre and has a high diversity of wild and cultivated grapevines. Archaeological findings indicate that winemaking activities have existed in Armenia for more than 6,000 years, viticulture being one of the most important activities of the modern Armenian agricultural sector. Despite this relevance, some grapevines in local collections have not yet been properly identified, thus hindering the efficient conservation, characterisation and eventual use of autochthonous genetic resources. In the present study, a combined SNP and SSR profiling strategy was used for the genetic identification of a series of grapevine accessions from the Grape Collection of the International Academy of Viticulture and Winemaking in Nalbandyan, presumed to be autochthonous Armenian varieties. The results provided useful information for the correct identification of these genetic resources, revealing multiple cases of synonyms, homonyms and misnames. The genetic data made it possible to confirm the pedigree proposed for some of the cultivars identified in this study and to clarify the origin of others. In addition, we propose, for the first time, a series of new trios and duos involving autochthonous Armenian grapevines. The singularity of this genetic pool compared to other Western and Central European varieties, as well as the potential novel sources of variability in traits of interest (e.g., seedlessness) that were found, highlight the importance of improving knowledge of the Armenian grapevine genetic pool.
... comm.). However, we cannot exclude that either natural or intentional selection took place in the region last two centuries [56,57], when the pressure of E. necator on grapevine cultivation became evident. Such a selection could explain the high frequency of resistance haplotype 149-208 within the Caucasian V. vinifera. ...
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Background Vitis vinifera L. is the most cultivated grapevine species worldwide. Erysiphe necator Sch., the causal agent of grape powdery mildew, is one of the main pathogens affecting viticulture. V. vinifera has little or no genetic resistances against E. necator and the grape industry is highly dependent on agrochemicals. Some Caucasian V. vinifera accessions have been reported to be resistant to E. necator and to have no genetic relationships to known sources of resistance to powdery mildew. The main purpose of this work was the study and mapping of the resistance to E. necator in the Caucasian grapes ‘Shavtsitska’ and ‘Tskhvedianis tetra’. Results The Caucasian varieties ‘Shavtsitska’ and ‘Tskhvedianis tetra’ showed a strong partial resistance to E. necator which segregated in two cross populations: the resistant genotypes delayed and limited the pathogen mycelium growth, sporulation intensity and number of conidia generated. A total of 184 seedlings of ‘Shavtsitska’ x ‘Glera’ population were genotyped through the Genotyping by Sequencing (GBS) technology and two high-density linkage maps were developed for the cross parents. The QTL analysis revealed a major resistance locus, explaining up to 80.15% of the phenotypic variance, on ‘Shavtsitska’ linkage group 13, which was associated with a reduced pathogen infection as well as an enhanced plant necrotic response. The genotyping of 105 Caucasian accessions with SSR markers flanking the QTL revealed that the resistant haplotype of ‘Shavtsitska’ was shared by ‘Tskhvedianis tetra’ and a total of 25 Caucasian grape varieties, suggesting a widespread presence of this resistance in the surveyed germplasm. The uncovered QTL was mapped in the region where the Ren1 locus of resistance to E. necator , identified in the V. vinifera ‘Kishmish vatkana’ and related grapes of Central Asia, is located. The genetic analysis conducted revealed that the Caucasian grapes in this study exhibit a resistant haplotype different from that of Central Asian grape accessions. Conclusions The QTL isolated in ‘Shavtsitska’ and present in the Caucasian V. vinifera varieties could be a new candidate gene of resistance to E. necator to use in breeding programmes. It co-localizes with the Ren1 locus but shows a different haplotype from that of grapevines of Central Asia. We therefore consider that the Caucasian resistance locus, named Ren1.2 , contains a member of a cluster of R-genes, of which the region is rich, and to be linked with, or possibly allelic, to Ren1 .
... Although the domestication centre of the grapevine is a long-contemplated subject, there is a consensus among archeologists and historians suggesting that the grapevine was distributed from Transcaucasia, which covers the borders of present-day Armenia, Georgia and Azerbaijan, together with north-eastern Anatolia. The region has been considered for a long time as the domestication centre for viticulture with the earliest examples of winemaking (McGovern, 2003;This et al., 2006;Maghradze et al., 2020). However, earlier, it was argued that the cultivation of vines for winemaking originated in the mountainous region between the Black Sea and the Caspian Sea, on the borders of the modern states of Turkey, Syria, Iraq, Iran and the former Soviet Union (Billiard, 1913;Johnson, 1989;Levadoux, 1956;Negrul, 1960;Ramishvili, 1983;Unwin, 1991). ...
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La “vite selvatica” (Vitis vinifera subsp. sylvestris) è considerata l’antenato selvatico dei moderni vitigni afferenti a Vitis vinifera. Sebbene attualmente questa entità sia ritenuta di scarsa valenza tassonomica, questo lessico botanico risulta spesso citato soprattutto nelle fonti letterarie storiche. In questo lavoro è stata effettuata, per la prima volta, una sintesi dei dati disponibili attraverso una minuziosa analisi delle fonti letterarie considerate pertinenti. I primi riferimenti alla “vite selvatica” sono stati rintracciati nell’Odissea di Omero, il poema epico composto probabilmente tra l’VIII ed il VII secolo a.C.
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Background Vitis vinifera L. is the most cultivated grapevine species worldwide. Erysiphe necator Sch., the causal agent of grape powdery mildew, is one of the main pathogens affecting viticulture. V. vinifera has little or no genetic resistance against E. necator and the grape industry is highly dependent on agrochemicals. Some Caucasian V. vinifera accessions have been reported to be resistant to E. necator and to have no genetic relationship to known sources of resistance to powdery mildew. The main purpose of this work was the study and mapping of the resistance to E. necator in the Caucasian grapes ‘Shavtsitska’ and ‘Tskhvedianis tetra’. Results The Caucasian varieties ‘Shavtsitska’ and ‘Tskhvedianis tetra’ showed a strong partial resistance to E. necator which segregated in two cross populations: the resistant genotypes delayed and limited the pathogen mycelium growth, sporulation intensity and number of conidia generated. A total of 184 seedlings of ‘Shavtsitska’ x ‘Glera’ population were genotyped through the Genotyping by Sequencing (GBS) technology and two high-density linkage maps were developed for the cross parents. The QTL analysis revealed a major resistance locus, explaining up to 80.15% of the phenotypic variance, on ‘Shavtsitska’ linkage group 13, which was associated with a reduced pathogen infection as well as an enhanced plant necrotic response. The genotyping of 105 Caucasian accessions with SSR markers flanking the QTL revealed that the resistant haplotype of ‘Shavtsitska’ was shared by ‘Tskhvedianis tetra’ and a total of 24 Caucasian grape varieties, suggesting a widespread presence of this resistance in the surveyed germplasm. The uncovered QTL was mapped in the region where the Ren1 gene of resistance to E. necator, identified in the V. vinifera ‘Kishmish vatkana’ and related grapes of Central Asia, is located. The genetic analysis conducted revealed that the Caucasian grapes in this study exhibit a resistant haplotype different from that of Central Asian grape accessions. Conclusions The QTL isolated in ‘Shavtsitska’ and present in the Caucasian V. vinifera varieties could be a new candidate gene of resistance to E. necator to use in breeding programmes. It co-localizes with the Ren1 locus but shows a different haplotype from that of grapevines of Central Asia. We therefore consider this gene, named Ren1-2, a member of a cluster of R genes, of which the region is rich, and linked with, or possibly allelic, to the Ren1.
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Grapevine (Vitis vinifera) is one of the most widely cultivated plant species of agricultural interest, and is extensively appreciated for its fruits and the wines made from its fruits. Considering the high socioeconomic impact of the wine sector all over the world, in recent years, there has been an increase in work aiming to investigate the biodiversity of grapevine germplasm available for breeding programs. Various studies have shed light on the genetic diversity characterizing the germplasm from the cradle of V. vinifera domestication in Georgia (South Caucasus). Georgian germplasm is placed in a distinct cluster from the European one and possesses a rich diversity for many different traits, including eno-carpological and phenological traits; resistance to pathogens, such as oomycetes and phytoplasmas; resistance to abiotic stresses, such as sunburn. The aim of this review is to assess the potential of Georgian cultivars as a source of useful traits for breeding programs. The unique genetic and phenotypic aspects of Georgian germplasm were unraveled, to better understand the diversity and quality of the genetic resources available to viticulturists, as valuable resources for the coming climate change scenario.
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A prospecting of habitats and mechanical support host species for the climber Eurasian wild grapevine, Vitis vinifera L. subsp. sylvestris (Gmelin) Hegi, was carried out on 13 natural populations situated along river bank forests, floodplains and colluvial positions in Georgia (Marneuli, Mtskheta and Gori districts, Gardabani Protected area and Lagodekhi Reserve), Armenia (Akhtala and Tavoush regions) and Azerbaijan (Quba region) during survey of 2013. The research demonstrated that Eurasian wild grapevine (Vitis vinifera subsp. sylvestris) is found in Southern Caucasus in a wide variety of habitats always linked to water availability. Punica granatum trees are the commonest mechanical support for wild grapevine in the South Caucasus and Hedera helix often shares the same support trees. However we documented wild grapevines climbing on other 24 different species of trees and large shrubs and, further, 32 associated species. We determined, four different clusters of localities using Structure software and the Weighted Neighbor Joining tree. These clusters are characterized by specific mechanical support and accompanying species. Other vines competing for host with Eurasian wild grapevine belong to the genera Clematis, Hedera, Humulus, Smilax and Vitis ssp.
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Background The mountainous region between the Caucasus and China is considered to be the center of domestication for grapevine. Despite the importance of Central Asia in the history of grape growing, information about the extent and distribution of grape genetic variation in this region is limited in comparison to wild and cultivated grapevines from around the Mediterranean basin. The principal goal of this work was to survey the genetic diversity and relationships among wild and cultivated grape germplasm from the Caucasus, Central Asia, and the Mediterranean basin collectively to understand gene flow, possible domestication events and adaptive introgression. ResultsA total of 1378 wild and cultivated grapevines collected around the Mediterranean basin and from Central Asia were tested with a set of 20 nuclear SSR markers. Genetic data were analyzed (Cluster analysis, Principal Coordinate Analysis and STRUCTURE) to identify groups, and the results were validated by Nei’s genetic distance, pairwise FST analysis and assignment tests. All of these analyses identified three genetic groups: G1, wild accessions from Croatia, France, Italy and Spain; G2, wild accessions from Armenia, Azerbaijan and Georgia; and G3, cultivars from Spain, France, Italy, Georgia, Iran, Pakistan and Turkmenistan, which included a small group of wild accessions from Georgia and Croatia. Wild accessions from Georgia clustered with cultivated grape from the same area (proles pontica), but also with Western Europe (proles occidentalis), supporting Georgia as the ancient center of grapevine domestication. In addition, cluster analysis indicated that Western European wild grapes grouped with cultivated grapes from the same area, suggesting that the cultivated proles occidentalis contributed more to the early development of wine grapes than the wild vines from Eastern Europe. Conclusions The analysis of genetic relationships among the tested genotypes provided evidence of genetic relationships between wild and cultivated accessions in the Mediterranean basin and Central Asia. The genetic structure indicated a considerable amount of gene flow, which limited the differentiation between the two subspecies. The results also indicated that grapes with mixed ancestry occur in the regions where wild grapevines were domesticated.
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