Euphytica (2012) 184:369–376
DOI 10.1007/s10681-011-0597-5
The plant size and the spine characteristics of the first
generation progeny obtained through the cross-pollination
of different genotypes of Cactaceae
Lucica Mihalte • Radu E. Sestras
Received: 5 June 2011 / Accepted: 29 November 2011 / Published online: 7 December 2011
Ó Springer Science+Business Media B.V. 2011
Abstract The main characteristics (plant diameter,
number of spine/areoles and length of spines) of 16
cactus hybrids were analysed and the genetic variabil-
ity and broad-sense heritability was studied. The
descendants were obtained through a cyclic cross-
pollination pattern, with the parental forms chosen
based on aesthetic considerations. Cross-pollination
among Rebutia senilis 9 Aylostera muscula, Rebutia
tarvitaensis 9 Aylostera muscula, Aylostera flavi-
styla 9 Rebutia senilis, Rebutia senilis 9 Aylostera
flavistyla, Aylostera muscula 9 Aylostera albiflora
and Rebutia senilis 9 Aylostera albiflora did not
succeed, whereas all of the other hand-pollinated
crosses succeeded and produced viable seeds. The
highest values of the analysed characters were
observed in the progeny of A. fiebrigii var. densise-
ta 9 R. senilis and A. buiningiana 9 A. vallegarden-
sis and the artificial selection to identify plants with
special decorative traits was extremely efficient among
them. In the F1 population of the studied crosses, a
large genetic diversity was found within hybrid
combinations (families), between combinations and a
different variation was recorded among the analysed
L. Mihalte (&)
R. E. Sestras
Department of Genetics and Plant Breeding,
University of Agricultural Science and Veterinary
Medicine, 3-5 Manastur, Cluj-Napoca 400372,
Romania
e-mail: mihaltelucica@yahoo.com
R. E. Sestras
e-mail: rsestras@yahoo.co.uk
traits. The broad-sense heritability ranged between
0.909 (plant diameter) and 0.948 (spines length). All of
the characters analysed, in the present experience, have
a strong genetic determinism, being greatly influenced
by the genotype and to a lesser extent, by the
cultivation conditions (greenhouse).
Keywords Cactaceae
F1 hybrids

Interspecific hybridisation
Inter-generic
hybridisation
Genetic diversity
Introduction
The Cactaceae family (order Caryophyllales) includes
plants with perennial photosynthetic succulent stems,
spines produced on modified axillary buds (termed
areoles) and colourful flowers with numerous stamens.
The multiple uses and the ability of cacti to thrive in
the arid and semiarid environments that other species
are unable to tolerate make it of interest to breeders
and molecular biologist seeking to develop crops for
areas typically unsuitable for conventional agriculture
(Nobel 2002). A primary breeding goal should be the
expansion of the production area for cacti by devel-
oping cold-tolerant cultivars because the current high-
yielding fruit, vegetable and forage varieties do not
survive between temperatures of -5 and -8°C (Loik
and Nobel 1991; Parish and Felker 1995; Wang et al.
1998).
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The genetic diversity in cultivated cacti is limited
by the small number of progenitors and the loss of
genetic variation during cultivation (Nobel 2002) and
most of the domesticated cacti grown for fruit or
ornamental flowers apparently originated from a
relatively narrow germplasm base. In the case of
Eastern cactus, over 100 distinct clones have been
described, but Meier (1995) proposed that all are
probably the descendants of three plants (two Hatiora
gaertneri and one H. rosea).
Cacti are threatened by the loss and degradation of
their habitat and illegal collection (Oldfield 1997;
Boyle and Anderson 2002); the most important factor
causing the loss of biodiversity is intensive land use
(Zak et al. 2004). The ‘‘Red List’’ of threatened species
of the International Union for the Conservation of
Nature (IUCN) provides a categorised list of species
based on their relative risk of extinction at a global
scale; the list includes at least 104 cacti species (7% of
all species) as vulnerable to extinction (IUCN 2008).
The majority of the studies regarding cacti hybrid-
isation have been restricted to the Opuntioideae
subfamily and little is known about the role of
hybridisation on Cactoideae, a more diverse subfam-
ily (Machado 2008). Therefore, in the present
research, we chose species from the Aylostera and
Rebutia genera and therefore, the aim of this study was
to identify potential genitors for breeding programs
and to improve the genetic diversity, which will permit
an efficient selection of new cultivars.
Materials and methods
Plant material and growth conditions
The Cactaceae species used in the present research were
grown in the greenhouse complex at the ‘‘Alexandru
Borza’’ Botanical Garden of Cluj-Napoca, Romania.
This Cactaceae collection comprises more than 4100
specimens belonging to 115 genera from the 241 total
known according to the Backeberg system (1968–1977).
Based on aesthetic considerations (shape of plant,
flower diameter and colour of flowers), we chose 19
genotypes as the genitors for the interspecific and
inter-generic hybridisations (Table 1).

Table 1 The main traits of the parental genotypes

Species Plant shapea No. spines/areoles Flower Flower
and length of spines diameter (cm) colours

Aylostera muscula Oval 23/2.4 mm 3.5 Orange

Aylostera flavistyla Globular 15–22/5–10 mm 3 Red
Aylostera fiebrigii Globular 30/40.1 cm 3.5 Orange
Aylostera fiebrigii var. densiseta Globular 40/50.1 cm 3.5 Orange
Aylostera spinosissima Globular 20–30/1.5 cm 3 Red
Aylostera narvacensis Globular 10–20/2–3 mm 4 Pink
Aylostera archibuingiana Globular 12/2–3 mm 4 Red
Aylostera buingiana Globular 14–16/0.6–1 cm 3 Pink
Aylostera vallegardensis Globular 20/3 cm 3.5 Pink
Aylostera albiflora Globular 15/5 mm 2.5 White
Rebutia senilis Globular 25/over 3 cm 3.5 Red
Rebutia senilis var. hyalacantha Globular 25/2 cm 3 Red
Rebutia travitaensis Globular 7–9/2–5 mm 6 Red
Rebutia cajasensis Globular 13/15.5 mm 3.5 Red
Rebutia spegazziana Cylindrical 23/4 mm 4 Dark red
Rebutia pseudominuta var schumaniana Globular 27/1.2 cm 2.5 Orange
Rebutia kuperiana Globular 13/15.5 mm 3.5 Red
Rebutia kuperiana var. spiniflora Globular 13/15.8 mm 3 Red
Rebutia donaldiana Globular 25/3 cm 3.5 Pink
a
All of the values represented the arithmetic mean

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Euphytica (2012) 184:369–376
The hybrid seeds were sown in small pots in a
mixture, composed of three parts of sterilised fibrous
soil, one part peat moss and one part washed sand
(Hewitt 1993) and were grown in the Botanical
Garden Cluj-Napoca, Romania. The seeds were free
of chemicals and fertiliser. In December, the minimum
temperature dropped to 5°C and the maximum
temperature reached 23.48°C in July.
Experimental design and statistical analyses
Using a cyclic hybridisation pattern, 14 interspecific
combinations (between two species of the same genus)
and 16 inter-generic combinations (between two plants
belonging to two different genera) was achieved.
To isolate a cross-pollinated flower from the
possibility of contamination by the pollen from other
plants and to distinguish the differences between
hybrid and non-hybrid fruits, each flower was insu-
lated individually with sticky paper and different
colours of thread were used for each flower (Mon-
dragon-Jacobo and Bordelon 1996).
A total of ten plants were pollinated for each
combination and every plant contained approximately
ten flowers. Hybrid seeds were obtained from 12
interspecific and 12 inter-generic combinations and
ten seeds were sown/pot. Viable F1 progeny were
obtained in 16 hybrid families and the plants were
analysed in the first year of vegetative growth (Fig. 1).
The low proportion of seed germination can be
explained by the different growth forms of cacti:
columnar cacti have a higher rate of seed germination
than globular cacti (Ortega-Baes et al. 2010). Aylos-
tera and Rebutia species are globular cacti and it is
Fig. 1 The first generation of the progeny of the studied cacti species at several weeks (a) and during the first year of vegetative
growth (b)
371
also possible that their seeds experience a dormancy
period (Mihalte et al. 2010).
The experiment was structured following a com-
pletely randomised (CR) design with three replica-
tions for each analysed trait (plant diameter, number of
spine/areoles and length of spines). The differences in
the characteristics of the progeny were analysed
statistically using ANOVA, where the mean of the
experiment was the control and the coefficient of
variability (CV%) for traits was computed. The data
are presented as the means and standard deviations for
each cyclic hybridisation.
Genetic parameters
The main characteristics of the F1 hybrids inform the
interspecific and inter-generic hybridisation were
calculated and are discussed in terms of the coeffi-
cients of variability within each hybrid family. The
uniformity of the traits was computed based on the
following coefficients: CV% \ 10%-low variability;
10% \ CV% \ 20%-medium variability and CV%
[ 20%-high variability (Reed et al. 2002; Sestras et al.
2009).
To determine the genetic proportion in the pheno-
typic expression of the studied characteristics, the
coefficient of heritability was also calculated, where
the individual of the F1 of a cross was the observa-
tional unit. The broad-sense heritability was noted and
computed by a classical model, H2 = rG2/rP2, where
rG2 is the genotypic variance and rP2 is the pheno-
typic variance (Holland et al. 2003; Piepho and
Möhring 2007; Sestras et al. 2008).
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Results muscula 9 Aylostera albiflora, Rebutia senilis 9 Ay-

The cross-pollination of parental forms
The cross-pollinations among Rebutia senilis 9
Aylostera muscula, Rebutia tarvitaensis 9 Aylostera
muscula, Aylostera flavistyla 9 Rebutia senilis,
Rebutia senilis 9 Aylostera flavistyla, Aylostera
lostera albiflora were unsuccessful (Table 2). All of
the other artificial hybridisations were successful and
produced viable seeds; however, a viable F1 genera-
tion was obtained in only 16 hybrid combinations. The
parental genotype, Aylostera fiebrigii, has been
reported to present a small capacity of germination
when it has been used for studying the germination

Table 2 Hybrid combinations and results of hybridisations

No. Mother genotype ($) Father genotype (#) Results of hybridization Viable F1
(fruits and seeds) descendants

Interspecific hybridisations
1. R. spegazziana R. senilis ? -
2. R. pseudodeminuta var. schumaniana R. senilis ? -
3. R. cajasensis R. senilis ? ?
4. R. senilis R. kupperiana var. spiniflora ? ?
5. R. senilis R. spegazziana ? ?
6. R. senilis var. hyalacantha R. spegazziana ? -
7. A. flavistyla A. archibuinigiana ? -
8. A. buiningiana A. vallegardensis ? ?
9. A. muscula A. vallegardensis ? ?
10. A. fiebrigii A. vallegardensis ? ?
11. A. muscula A. buiningiana ? ?
12. A. buinigiana A. flavistyla ? -
13. A. muscula A. albiflora - -
14. A. spinosissima A. albiflora ? ?
Inter-generic hybridisations
15. R. senilis A. muscula - -
16. R. tarvitaensis A. muscula - -
17. R. cajasensis A. muscula ? ?
18. A. flavistyla R. senilis - -
19. A. fiebrigii var. densiseta R. senilis ? ?
20. A. spinosissima R. senilis ? -
21. A. muscula R. kupperiana var. spiniflora ? ?
22. A. narvaecensis R. kupperiana var. spiniflora ? ?
23. A. spinosissima R. spegazziana ? -
24. R. cajasensis A. archibuinigiana ? ?
25. R. senilis A. archibuinigiana ? ?
26. R. donaldiana A. buiningiana ? ?
27. R. kupperiana var. spiniflora A. buiningiana ? -
28. R. cajasensis A. flavistyla ? ?
29. R. senilis A. flavistyla - -
30. R. senilis A. albiflora - -
‘‘?’’ means that the specified hybrid combinations produced fruits and seeds, respectively and viable F1 progeny; ‘‘-’’ means that the
specified hybrid combinations did not produce fruits and seeds, respectively, or viable F1 progeny

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percentages of different cacti species (Mihalte et al.
2009).
The plant diameter
The plant diameters of the progeny were variable
(Table 3), with the average of the F1 hybrids in the
studied families varying significantly compared with
the control.
High values of plant diameter, suggesting high
vigour, presented in the hybrids of the following
combinations: R. senilis 9 R. kupperiana var. spini-
florum, A. buiningiana 9 A. vallegardensis, A. mus-
cula 9 A. vallegardensis. In general, the hybrids
derived from the interspecific combinations were
more vigorous than the hybrids from the inter-generic
combinations.
The following different variations of characteristics
in the F1 populations were noted: a low variability
among the hybrids of R. cajasensis 9 R. senilis and
R. senilis 9 R. kupperiana var. spiniflorum; a large
variability for the A. narvaecensis 9 R. kupperiana
var. spiniflorum and R. senilis 9 A. archibuinigiana
hybrid family and a medium variability for the
remainder of the hybrid combinations.
The number of spines/areoles
The variability of the number of the spines/areoles in
the F1 hybrids was quite small; only the combinations
A. buiningiana 9 A. vallegardensis and R. cajasen-
sis 9 A. archibuinigiana presented hybrids with a
medium variability of this trait (Table 3).
Compared with the mean of experience (21.0), the
number of spines/areoles in the F1 hybrids showed
positive values for the following crosses: A. fiebrigii var.
densiseta 9 R. senilis, A. muscula 9 R. kupperiana
var. spiniflorum, A. muscula 9 A. vallegardensis and A.
spinosissima 9 A. albiflora. Of all of the hybrid
combinations, the lowest number of spines/areoles

Table 3 The plant diameter, the number of spines/areolas and the length of spines, coefficient of variability and significance of
values* of the F1 hybrids
No. Hybrid combination Plant diameter CV% No. of spines/ CV% The length CV%
($ 9 #) (cm) areolas of spines (mm)

1. R. cajasensis 9 R. senilis 2.6 ± 0.1 8.4 14.5 ± 6.2°°° 8.9 20.0 ± 4.2* 11.3
2. R. senilis 9 R. kupperiana var. spiniflorum 3.2 ± 0.7** 8.4 22.3 ± 1.6 7.7 15.8 ± 0.1 7.2
3. R. senilis 9 R. spegazziana 1.8 ± 0.8°° 11.9 22.5 ± 1.8 5.7 17.2 ± 1.4 9.8
4. A. buiningiana 9 A. vallegardensis 3.3 ± 0.8** 23.0 15.5 ± 5.2°°° 13.4 20.6 ± 4.8* 9.4
5. A. muscula 9 A. vallegardensis 3.2 ± 0.7** 13.3 23.5 ± 2.8** 5.5 9.6 ± 6.3°° 17.7
6. A. fiebrigii 9 A. vallegardensis 2.5 ± 0.0 19.0 20.5 ± 0.2 6.3 26.2 ± 10.4*** 13.8
7. A. muscula 9 A. buiningiana 2.6 ± 0.1 8.6 22.5 ± 1.8 5.7 9.8 ± 6.0°° 13.6
8. A. spinosissima 9 A. albiflora 3.6 ± 1.1*** 8.2 23.3 ± 2.6** 11.8 15.5 ± 0.3 13.6
9. R. cajasensis 9 A. muscula 1.5 ± 1.0°°° 14.4 16.8 ± 4.0°°° 10.2 8.8 ± 7.1°°° 20.9
10. A. fiebrigii var. densiseta 9 R. senilis 2.4 ± 0.2 11.3 27.8 ± 7.1*** 6.2 27.1 ± 11.3*** 11.3
11. A. muscula 9 R. kupperiana var. spiniflorum 2.8 ± 0.3 6.2 25.5 ± 4.8*** 5.1 11.4 ± 4.4° 17.8
12. A. narvaecensis 9 R. kupperiana var. 2.2 ± 0.3 24.7 21.5 ± 0.8 6.0 12.8 ± 3.0 23.8
spiniflorum
13. R. cajasensis 9 A. archibuinigiana 2.1 ± 0.4 12.3 17.5 ± 3.2°°° 11.9 8.5 ± 7.4°°° 37.5
14. R. senilis 9 A. archibuinigiana 1.9 ± 0.6° 28.2 19.8 ± 0.9 8.6 14.4 ± 1.4 18.7
15. R. donaldiana 9 A. buiningiana 2.4 ± 0.1 17.7 20.8 ± 0.1 10.7 18.2 ± 2.4 28.8
16. R. cajasensis 9 A. flavistyla 2.0 ± 0.5° 12.7 21.5 ± 0.8 8.9 17.6 ± 1.8 19.0
Mean of experiment (control) 2.5 14.2 21.0 8.3 15.8 17.1
LSD 5% 0.5 2.5 3.7
LSD 1% 0.7 3.3 4.9
LSD 0.1% 0.9 4.3 6.4
*, **, ***/°, °°, °°° Significant at P \ 0.05, 0.01 and 0.001 (* means positive and ° means negative, respectively)

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was recorded for the hybrids derived from R. cajasensis
9 R. senilis and A. buiningiana 9 A. vallegardensis.
Regarding the genitors, the smallest number of spine/
areoles was recorded in R. cajasensis and A. archibuin-
igiana species and it appears that this characteristic was
transmitted to the F1 progeny derived from these
crosses.
The length of spines
Very small spines presented the hybrids of the
following crosses: R. cajasensis 9 A. muscula, R.
cajasensis 9 A. archibuinigiana and A. muscula 9 A.
vallegardensis (Table 3).
The results showed that when R. cajasensis and A.
archibuinigiana were used as the genitors, the length of
the spines was transmitted to the progeny with sufficient
fidelity and the hybrids progeny inherited the small
number of spines/areoles. A. fiebrigii and A. vallegard-
ensis, both having longer spines than most of the studied
species, passed this trait to their descendants.
The variability and the heritability of the analysed
traits
The obtained data showed the existence of a large
diversity, where the resulting variability was mani-
fested within the hybrid combinations (families) and
between the combinations. We also found a different
variation based on the characteristic that was analysed
(Fig. 2).
The lowest coefficient of variability was recorded
for the number of spines/areoles (18.0%), whereas a
wide variability was recorded for the plant diameter
(26.9%) and length of spines (38.4%). Therefore, the
CV% values indicate that it is possible to identify
Length of the
0.948
22.40
spines
Number of 0.943
spines/areolas 18.00
0.909
Plant diameter
26.90
0.00 5.00 10.00 15.00 20.00 25.00
The variability The broad-sense heritability
Fig. 2 The variability and the broad-sense heritability of the
studied characteristics in the first-generation progeny (F1)
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Euphytica (2012) 184:369–376
hybrids with small plant diameters and small lengths
of spines and with large plant diameters and large
lengths of spines in any combination (both forms are
suitable for new ornamental forms).
The broad-sense heritability (Fig. 2) ranged
between 0.909 (plant diameter) and 0.948 (spine
length). All of the characters analysed, at least in terms
of this experience, have a strong genetic determinism,
being strongly influenced by genotype and to a lesser
degree, by the culture medium (greenhouse).
Discussion
The ability of pollen to germinate is an important
aspect in cross-pollination (Oyiga et al. 2010). In
previous studies, the pollen germination ability of the
parental plants used in this study was investigated and
detected as being high (Mihalte et al. 2010). Even with
a high percentage of viable pollen, the cross-pollina-
tion between Aylostera flavistyla and Rebutia senilis
did not succeed with using Aylostera flavistyla as
either the male or female parent. Thus, Aylostera
flavistyla can be considered incompatible, as can
Hatiora (Boyle et al. 1994) and Schlumbergera (Boyle
1997). In addition, manual cross-pollination using
wild Stenocereus stellatus failed between some
domesticated and wild genotypes, suggesting a pollen
incompatibility (Casas et al. 1999). The hybridisation
experiments by Hentzschel and Hentzschel (2001) of
Sulcorebutia, Weingartia, Rebutia and Echinopsis
have produced the following results: most of the
hybridisation failed between Sulcorebutia and Rebutia
padcayensis, even though the latter closely resembles
Sulcorebutia and with several other Rebutia species.
For other genera (Selenicereus and Hylocereus), inter-
generic crosses appear to be a successful approach to
increase diversity. Tel-Zur et al. (2003) obtained
triploid, pentaploid, hexaploid and aneuploid hybrids
in inter-generic crosses (Selenicereus and Hylocere-
us). However, Lichtenzveig et al. (2000) reported
viable hybrids from inter-generic and interspecific
crosses between Selenicereus and Hylocereus.
High vigour, as observed for R. senilis 9 R.
kupperiana var. spiniflorumand and A. buiningi-
30.00 ana 9 A. vallegardensis, is of considerable impor-
tance to breeders, especially for the Rebutia genus,
which are generally small (Pilbeam 1997). For breed-
ers, vigour correlates with increased photosynthetic
Euphytica (2012) 184:369–376
area and improved root anchorage (Mondragon-
Jacobo and Bordelon 1996).
McIntosh (2002) correlated vigour with the size of
the plants for species of Ferocactus and showed that
large plants grow more than small plants. However, in
terms of percent change in volume, the relationship
was the opposite: large plants grew less as a percent of
size than small plants. In addition, the number of
flowers produced by a plant was significantly and
positively correlated with the size of the plant, but the
flower abortion rates (Burd 1998) and the number of
seeds per fruit were not influenced by this factor.
The study of the seminal progeny obtained from
hand-pollination showed that central spines lack in F1
hybrids, as they do in their parents. For most crop
cacti, the presence of spines is an inconvenience and
significant efforts to develop new cultivars are focused
on spineless cultivars (Mondragon-Jacobo and Bord-
elon 1996). For ornamental cacti, as are the species
described here, the presence of spines is a positive
element for their decorative traits. Peharec et al.
(2010) evaluated the spines of pot-grown Mammillar-
ia plants and in vitro-regenerated shoots and found
that the pot-grown plants had 16–17 spines per areole,
whereas the in vitro-grown shoots were normal but
displayed a lower number of spines (11–12). Parra
et al. (2010) argued that genetic diversity should be
conserved and should be promoted by artificial
selection, which, despite the high levels of gene flow,
moderate the genetic structure between wild and
managed populations.
A. fiebrigii var. densiseta 9 R. senilis, A. mus-
cula 9 R. kupperiana var. spiniflorum and A. mus-
cula 9 A. vallegardensis presented a large number of
spines/areoles. The obtained results suggest the poly-
genic inheritance of this trait, whereas the genitors of
these inter-generic and interspecific combinations
were noted for their large number of spines/areoles.
With regard to breeding programmes, particular
emphasis should be placed on the mode of inheritance
of basic traits, such as the length of spines, number of
spines/areoles, fruit colour, size and shape (Mondrag-
on-Jacobo and Bordelon 1996).
The highest values of the analysed characteristics
were noted for A. fiebrigii var. densiseta 9 R. senilis
and A. buiningiana 9 A. vallegardensis and the
artificial selection to identify plants with special
decorative traits was extremely effective for these
combinations. It is possible that these results may be
375
due mainly to the influence of the female parent, but
we do not exclude the possible effects of the male
genitor and of course, those caused by the environ-
mental conditions, culture and interactions between
the genotype and ecotype.
The high genetic variability shown in the present
research allows an efficient selection, but further
investigation should be performed by examining the
levels of allozyme variation and the genetic structure
(Figueredo et al. 2010).
For most cross-pollinations, as was the case of the
cactus species described here, the efficiency of
phenotypic selection is decisively affected by the size
of the heritability coefficient (Leite et al. 2006). Thus,
the phenotypic selection for all of the studied charac-
ters in the populations represented by the F1 hybrids
can be efficient.
Because the inheritance from genitors and the fixing
of such characteristics was possible in the F1 hybrid
descendants, studies of cacti germplasm to perform
artificial variability followed by artificial selection are
absolutely justified and extremely useful to obtain new
varieties. The obtained results are encouraging find-
ings, which indicate that the studied characteristics are,
overall, under strong genetic control and that a
genotype with a given performance could be selected
and genetically improved. The narrow-sense heritabil-
ity and the additive genetic variance have not been
studied in this experience because these aspects are
inappropriate for hybrids, owing to the genetic dis-
equilibrium inherent in their origin (Gordon 1999).
The obtained results indicate that through artificial
hybridisation under a suitable choice of genitors,
progeny with major features for selection that have
genetically improved ornamental qualities can be
produced, as was found for the crosses between A.
fiebrigii var. densiseta 9 R. senilis and A. buiningi-
ana 9 A. vallegardensis.
Acknowledgments This study was financed by the POSDRU/
89/1.5/S/62371 project (,,Postdoctoral School of Agriculture
and Veterinary Medicine’’, Romania).
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