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EFFECT OF NICKEL ON SEED GERMINATION OF ALYSSUM SPECIES WITH

POTENTIAL FOR PHYTOMIMING IN ALBANIA

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© by PSP Volume 27 – No. 3/2018 pages 1345-1352
EFFECT OF NICKEL ON SEED GERMINATION OF
ALYSSUM SPECIES WITH POTENTIAL FOR
PHYTOMIMING IN ALBANIA
Dolja Pavlova1,*, Donaltina Vila2, Kalina Vila2, Aida Bani2, Besmira Xhaferri2
1
Department of Botany, Faculty of Biology, University of Sofia, Sofia, Bulgaria
2
Agro-Environmental Department, Faculty of Agronomy and Environment, Agricultural University of Tirana, Koder-Kamez,
Tirana, Albania
ABSTRACT
The aim of this work was to compare the effect
of Ni on seeds of Alyssum markgrafii and A. murale,
an obligate and a facultative Ni hyperaccumularors
respectively, distributed on the Balkans. Seeds col-
lected from natural populations of the species in Al-
bania were germinated with standard solutions of
0.5, 1, 2, 4, 6, 8 mM Ni as NiCl2 6 H2O in distilled
water and compared with germinated seeds in dis-
tilled water. The results showed that Ni influenced
the germination process and differences were found
not only between the species but also between the
populations. The germinability decreased with ele-
vation of Ni concentrations. The relative frequency
of germination in A. markgrafii was more spread out
through time. The seeds of A. murale displayed
greater homogeneity and higher synchrony of germi-
nation. The seeds of A. murale were less sensitive (or
tolerant to) when treated with higher metal concen-
trations and germination was less influenced com-
pared to A. markgrafii. Lower concentrations of Ni
stimulated hypocotyl elongation but inhibited root
elongation in both species. The roots were more sen-
sitive to Ni compared to the hypocotyl. The abnor-
malities at early stage of seed germination found in
both species disturbed the normal growth of the
seedlings.
KEYWORDS:
Alyssum, nickel, hyperaccumulators, phytotoxicity, seed
germination
INTRODUCTION
The Balkan Peninsula is a region with numer-
ous obligate and facultative Alyssum hyperaccumu-
lator species of potential utility for extracting Ni
from soil [1, 2, 3]. Suitable species must be relatively
easy to collect as bulk seed accessions and have high
success rates of germination, establishment and
growth [4].
Nickel is a highly mobile trace metal that tends
to accumulate in newly formed plant organs and
1345
Fresenius Environmental Bulletin
seeds [5]. The information on the presence and Ni
localization in seeds is important for better under-
standing of plant reproductive strategies and seed
germination. Nickel localization patterns have been
determined for ten Alyssum Ni-hyperaccumulator
species/ecotypes [6] but data for the localization of
Ni in seeds are scarce. Munzuroglu and Geckil [7]
considered the effects of metal on germination of
seeds depend on differences in seed structure and
particularly seed coats. Nickel distribution in seeds
and seedlings during germination studied in Ni hy-
peraccumulator Berkheya coddii was demonstrated
by Groeber et al. [8].
High Ni levels exert toxic effects on seed ger-
mination and information on the seed germination
contributes to better understanding of plant adapta-
tion to habitats but literature investigating the same
on hyperaccumulators is rather scarce. The few pub-
lished works are restricted to the germinability as-
sessment [9, 10, 11]. Seed germination is one of the
most important stages in crop development that pre-
determines establishment of healthy crop, better
growth and yield [12, 13, 14]. The seed germination
and the root elongation tests are the most appropriate
and commonly used to prove the phytotoxicity in
vascular plants [15, 16, 17].
The work presented in this paper aims to com-
pare the effect of Ni phytotoxicity with different con-
centrations on seed germination, root and hypocotyl
elongation in two Ni hyperaccumulator species (A.
markgrafii O.E. Schultz and A. murale Waldst. &
Kit.), both of economic importance in Albania.
MATERIALS AND METHODS
Study material and sites. The plant material
was collected from Ni-hyperaccumulator species A.
murale and A. markgrafii distributed on serpentine
terrains in Albania. Seed material from A. murale
was collected from two localities: Prrenjas-Do-
mosdova (P) 41°00′08″N, 20°33′11″E (400 m a.s.l.)
and Pojske-Pogradec (Po) 40°59′55,″ N 20°38′09″E
(700 m a.s.l.) in 2014 and 2015, respectively. Seeds
from the serpentine endemic species A. markgrafii
were collected from Gjegjan (G), 41°55′47″N
© by PSP Volume 27 – No. 3/2018 pages 1345-1352
20°00′09″E (700 m a.s.l.) and from Kukës (K),
42°00'35"N, 20°17'44"E (1300 m a.s.l.) in 2014. The
soil chemical composition of these sites was previ-
ously analyzed [1, 18] and reflected their ultramafic
origin, i.e. between 1100-2150 mg kg-1 of Cr, 1080-
3100 mg kg-1 of Ni and 104 -280 mg kg-1 of Co. Seed
material was collected from 50 individuals per pop-
ulation. The seeds were extracted manually from
fruits and those of bad quality (damaged, empty and
immature) were rejected. The received seeds from A.
markgrafii were low in comparison to A. murale,
probably due to lower seed production. The material
was collected and stored according to international
guidelines [19]. Vouchers were deposited in the Ag-
ricultural University of Tirana.
Seeds germination, root and hypocotyl elon-
gation. Seed germination abilities were evaluated in
laboratory conditions following the method of Mun-
zuroglu and Geckil [7]. According to the authors this
method reduces the effect of counteranions associ-
ated with other metals or other metal cations nor-
mally present in the soil. Standard solutions of 0.5,
1, 2, 4, 6, 8 mM Ni as NiCl2 6H20 were prepared with
250 ml distilled water. The range of concentrations
is based on metal concentrations commonly encoun-
tered in non-serpentine soil to their several-fold
higher concentrations measured in the serpentine
soils. Distilled water was used as a control. The
seeds were surface-sterilized in 2% (w/v) calcium
hypochlorite solution for 10 min, then washed and
soaked for 24 h in distilled water before treatment
following Ghaderian et al. [11]. Harvested seeds of
A. murale and A. markgrafii were tested for their ger-
minability without exposing them to any dormancy-
relieving treatments. For each treatment 50 seeds
from A. murale were used while from A. markgrafii
they were 30 seeds because of lack of enough mate-
rial received. The seeds were germinated in sterile 9
cm Petri dishes on filter paper moistened with Ni so-
lutions following Munzuroglu and Geckil [7]. Seed
incubation was in a temperature-controlled growth
cabinet set at 21/21o C (day/night cycle). Petri dishes
were exposed to a 12 h light/12 h dark cycle. The
illumination was provided by white fluorescent
tubes with mean photon flux density of 220 mmol-2
m-1 s-1 at seed level. Roots were considered germi-
nated when 1 mm radicle has appeared. The number
of seeds germinated was counted daily till the end of
the experiment (three consequent days with constant
germinated seeds). Seed root and hypocotyl length
were measured using stereomicroscope STEMI
2000-C, Zeiss. The germinability (G%), mean ger-
mination time (MT), coefficient of variation of the
germination time (CVt) and mean germination rate
(MR), uncertainty of germination (U), and synchro-
nization index of germination (Z) were used to de-
scribe germination process for both species and their
studied populations following Ranal and Santana
1346
Fresenius Environmental Bulletin
[20] and Ranal et al. [21]. All treatements were per-
formed with three replications.
Data analysis. The data were analysed with
one-way analysis of variance (ANOVA) to deter-
mine which treatments differed from each other
when germinability, root and hypocotyl elongation
were used as variables. The data for both species and
their population were analysed separately. In cases
when the ANOVA showed significance of differ-
ences (P < 0.05), pairwise mean comparisons of var-
iables, a least significant difference (LSD) test was
performed. The statistical analysis was performed
using Microsoft Excel 2010.
RESULTS
Seeds germination. The germinability of both
Alyssum species was significantly different (Table 1)
and the results demonstrated that Ni concentrations
influenced the germination process. In general, the
germinability decreased with elevation of the Ni
concentrations used for seed treatments for all stud-
ied populations. Alyssum murale demonstrated
higher germinability compared to A. markgrafii. The
percentage of germinated seeds in the controls for A.
murale populations (87.33% and 68%, respectively)
and for A. markgrafii populations (51.55% and 50%,
respectively) was higher than in Ni treatments. The
effective concentration causing ~50% inhibition of
germination was not reached during the experiment
with seeds in both populations of A. murale. This
concentration for A. markgrafii was 6 mM Ni for the
seeds from Kukës. There was a significant effect of
treatments on seed germination in both A. murale
populations (LSD = 7.62, P < 0.05 and LSD = 16.77,
P < 0.05, respectively). Significant differences were
found between the germinability of seeds from the
control and in some cases between seeds treated with
lower concentrations Ni (0.5 and 1 mM) (Table 1).
The concentration of 8 mM Ni decreased the germi-
nability in A. murale with 9% and 11%, respectively
for both populations. No significant differences in
germinability were shown between the seeds of A.
murale treated with 2, 4, 6 and 8 mM Ni. The results
of germinability of seeds in populations of A.
markgrafii demonstrated also significant effect of Ni
(LSD = 15.5, P < 0.05 and LSD = 16.11, P < 0.05
respectively). The highest Ni concentration de-
creased the germinability with 13% and 29%, re-
spectively for A. markgrafii.
The mean germination time (MT) was clearly
different between the species and populations (Table
1). It was higher for A. markgrafii where MT was
~11-12 days while for A. murale it was ~3 and ~5
days, respectively for both populations. The seeds of
A. murale started their germination ~2 days after
treatment while for A. markgrafii it was after 4-5
days. The relative frequency of germination in A.
© by PSP Volume 27 – No. 3/2018 pages 1345-1352 Fresenius Environmental Bulletin

markgrafii was more spread out through time, with a
peak of germination at ~12th and 14th days respec-
tively for both populations. The germination was
less spread out in time in A. murale and a strong ger-
mination peak occurred at the 4th day for seeds from
the first population and at the 6th day after beginning
of the experiment for the second population (Table
1).
The mean germination rate (MR) calculated as
the reciprocal of the mean germination time (MT)
was the highest for A. murale (Prrenjas) and lowest
for A. markgrafii (Kukës). The seeds of A. murale
presented greater homogeneity (lower value of CVt)
and higher synchrony of germination (lower value of
U and higher value of Z) compared to A. markgrafii.
The differences of the germination process were
found not only between the species but also between
their populations, higher between A. murale popula-
tions.
Root and hypocotyl elongation. The root and
hypocotyl elongation of seeds treated with Ni was
also affected by elevated concentrations in both spe-
cies (Fig. 1). In general, the length of root treated
with different Ni concentrations decreased compared
to the control. The length of the root after Ni appli-
cation was significantly different from the controls
in A. murale (LSD = 2.88, P < 0.05 and LSD = 2.10,
P < 0.05, respectively) and A. markgrafii (LSD =
4.39, P < 0.05 and LSD = 7.72, P < 0.05, respec-
tively) populations (Figs. 1A and C). The effect of
inhibition on the root growth was higher after treat-
ments with ≥ 2 mM Ni in both species. The concen-
tration of 1 mM Ni reduced root length about 50%
for A. murale and A. markgrafii from Gjegjan. For
the seeds of A. markgrafii (Kukës) this concentration
was 0.5 mM Ni. The root length was more or less
equal when the seeds were treated with 6 and 8 mM
Ni and no significant differences were found.

TABLE 1
The calculated values for germinability (G%), mean germination time MT (days), coefficient of variation
of the germination time (CVt%), mean germination rate (MR), uncertainty (U) and synchronization of
germination (Z) for A. murale and A. markgrafii populations. Values are the means of three replicates per
population ± SD (n=6). Different letters show significant differences based on one-way ANOVA and LSD
test (P < 0.05) performed separately for each population.

Species &
Ni
popula- G (%) MT (day) CVt (%) MR (1/day) U Z
(mM)
tions
0 87.33±1.15a 3.00±1.06a 33.95±15.47a 0.35±0.10a 1.04±0.13ab 0.63±0.05a
0,5 80.67±5.03ab 2.78±0.85ab 24.53±3.23a 0.37±0.09a 1.13±0.33a 0.52±0.15b
1 86±4a 2.97±0.92bc 24.47±1.18a 0.34±0.09a 1.36±0.50bc 0.45±0.18b
A. murale
2 82±2ab 3.28±0.82c 28.05±4.51a 0.31±0.07a 1.67±0.17cd 0.31±0.04bc
(P)
4 84±7.21ab 3.26±1.00c 31.74±11.28a 0.32±0.08a 1.64±0.23c 0.35±0.07bc
6 83±5.03ab 3.57±0.80c 31.55±4.67a 0.28±0.05a 1.95±0.18d 0.26±0.04c
8 78±4b 3.58±0.99c 30.27±6.12a 0.29±0.07a 1.85±0.25d 0.30±0.07bc
0 68±4a 4.15±0.14a 22.13±1.35a 0.23±0.05a 1.77±0.13ab 0.31±0.06a
0,5 64±11.13ab 4.52±0.29a 24.19±0.43a 0.21±0.01b 2±0.08ab 0.24±0.01bc
1 55±17.00abc 4.94±0.27a 20.96±4.14a 0.19±0.01bc 1.87±0.24ab 0.29±0.03ab
A. murale
2 51±10.06bc 5.15±0.42a 20.14±2.37a 0.18±0.01c 1.96±0.14ab 0.25±0.02bc
(Po)
4 55±1.15abc 5.07±0.25a 22.81±3.85a 0.19±0.01bc 2.00±0.13a 0.24±0.05bc
6 43±7.02c 5.27±0.48a 22.26±0.75a 0.18±0.02c 1.99±0.08b 0.23±0.02bc
8 57±9.86abc 5.3±0.1a 22.20±1.48a 0.18±0.05c 2.13±0.03b 0.21±0.01c
0 51±5.13a 10.52±0.75abc 27.65±1.05a 0.09±0.01ab 2.63±0.19ab 0.08±0a
0,5 48±5.03ab 9.59±0.82ab 41.69± 0.87ab 0.10±0.01a 2.86±0.51ab 0.09±0.06a
A. 1 49±5.54ac 9.77±0.77a 44.67±9.42b 0.09±0.01ab 2.75±0.27ab 0.08±0.02a
markgrafii 2 30±3d 11.61±0.40bc 38.43±4.93ab 0.08±0.00ab 2.61±0.20ab 0.04±0.02a
(G) 4 26.33±11.54d 10.75±2.18abc 29.19±4.54aa 0.09±0.02ab 2.19±0.61a 0.09±0.06a
6 34±4.04bcd 12.41±0.92c 32.32±5.41ab 0.07±0.01b 2.68±0.35b 0.08±0.04a
8 38.00±15.58bad 12.63±2.65c 31.24±18.29ab 0.07±0.02ab 2.63±0.48b 0.07±0.02a
0 50±5.77a 8.87±1.13a 38.87±9.76a 0.11±0.01a 2.62±0.50abc 0.12±0.07a
0,5 38.67±5.13ab 10±1a 32.05±6.29a 0.10±0.01ab 2.71±0.13abc 0.08±0.01ab
A. 1 45.67±14.01ab 9.90±1.26a 38.47±6.23a 0.09±0.01ab 2.99±0.31a 0.06±0.02ab
markgrafii 2 33±10b 11.27±0.45a 29.60±5.29a 0.08±0.01b 2.62±0.25abc 0.06±0.04ab
(K) 4 31±7.21b 11.64±2.12a 31.89±13.89a 0.08±0.01b 2.49±0.39b 0.01±0.04ab
6 21±13.32b 11±1.80a 32±8.07a 0.09±0.01b 2.15±1.01c 0.03±0.02b
8 21.00±8.54b 11.76±0.48a 34.11±13.26a 0.08±0.00b 2.00±0.10c 0.12±0.07ab

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© by PSP Volume 27 – No. 3/2018 pages 1345-1352 Fresenius Environmental Bulletin

FIGURE 1
Root and hypocotyl length in Alyssum murale (A and B) and Alyssum markgrafii (C and D) populations.
Values are the means of three replicates per population ± SD (n=6). Abbreviations are used to indicate
different populations: Prrenjas (P), Pogradec (Po), Gjegjan (G) and Kukës (K). Different letters show
significant differences based on one-way ANOVA and LSD test (P < 0.05) performed separately for
each population.

In general, the hypocotyl length in the control
was lower compared to the length of hypocotyl for
seeds treated with lower concentrations of Ni (0.5
and 1 mM) (Figs. 1B and D). After treatment with
such concentrations the length of the hypocotyl de-
creased and reached 50% inhibition for seeds treated
with 8 mM Ni compared to the control in both pop-
ulations of A. markgrafii. For A. murale 50% inhibi-
tion was not reached in both populations. The statis-
tical analysis and pairwise mean comparisons of var-
iables demonstrated significantly differences be-
tween the control and seeds treated with lower Ni
concentrations only in A. murale from Prrenjas (LSD
= 1.94, P < 0.05). In the population of A. murale from
Kukës the hypocotyl length in the control was sig-
nificantly different from seeds treated with higher
concentrations (LSD = 0.85, P < 0.05). For both pop-
ulations of A. markgrafii the hypocotyl length in the
control was significantly different from seeds treated
with higher concentrations (LSD = 2.08, P < 0.05
and LSD = 2.14, P < 0.05, respectively).
The effect of Ni on root and hypocotyl length
was different in both species and their populations.
Lower concentrations of Ni stimulated hypocotyl
growth but inhibited root elongation. Root seedlings
in both species and populations were more sensitive
to Ni compared to hypocotyl. Although the length of
the roots in the controls was higher compared to the
length of the hypocotyl, it decreased quickly and
reached lower values in samples treated with 6 and 8
mM Ni.
Nickel impact on seedlings. Seeds of both spe-
cies and populations treated with different Ni con-
centrations showed abnormalities in their germina-
tion, including chlorosis, necrosis, coiling of the root
and the hypocotyl, lack of hairs in the zone of root
hairs, a short zone of root elongation and the appear-
ance of root deviation very near to the root tip, brown
cotyledons and hypocotyl of the seedlings or three
cotyledons instead of a pair (Fig. 2). These abnor-
malities varied considerably for each Ni treatment.
Soon after germination in 0.5 mM Ni, the root and
hypocotyl started coiling (Figs. 2D and E) and brown
color of the hypocotyl and the cotyledons appeared
(Figs. 2G and H). The root coiling was more than
70% of the seedlings of A. murale treated with 0.5
mM Ni while for A. markgrafii it was more than
50%. The percentage of the root coiling increased
and varied between 90% and 100% when the seeds

1348
© by PSP Volume 27 – No. 3/2018 pages 1345-1352 Fresenius Environmental Bulletin

were treated with 8 mM Ni in both species and pop-
ulations. The percentage of the hypocotyl coiling (in
the area near to the cotyledons) was lower compared
to the root coiling in both species and all Ni treat-
ments. Frequently, cotyledons with curved margins
were found (Fig. 2F). The cotyledons demonstrated
also chlorosis and necrosis when Ni concentrations
were higher (Figs. 2B and C). The percentage of the
cotyledons with chlorosis was higher in A. murale.
Most often roots without hairs appeared in both spe-
cies when treated with Ni concentrations above 1
mM. The seed germination stopped soon after the
appearance of radical in higher Ni concentrations
(Fig. 2I). Also, a short zone of root elongation (Fig.
2J) and the appearance of root deviation very near to
the root tip (Fig. 2K) were found in seeds treated
with 6 and 8 mM Ni. Rarely, three cotyledons in-
stead of a pair were observed (Fig. 2L).

FIGURE 2
Different patterns of abnormally germinated seed of Alyssum species after Ni application. A: normally
germinated seedling; B-C: chlorosis and necrosis on the leaves; D-E: coiling root and hypocotyl; F: cot-
yledons with curved margins; G-H: brown color of hypocotyl and cotyledons; I: seed germination
stopped after the appearance of radical in higher Ni concentrations; J: short zone of root elongation;
K: root deviation very near to the root tip; L: appearance of three cotyledons instead of a pair. Scale in
millimeters (mm).

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© by PSP Volume 27 – No. 3/2018 pages 1345-1352
DISCUSSION
Both Alyssum species and populations showed
differences in the germination process, probably re-
lated to the species physiology. The germinability
was higher for the seeds of A. murale. The difference
in germinability between the two studied popula-
tions of A. murale was possibly in relation to the seed
age. In previous studies on dormancy and germina-
tion percentage of Alyssum bertolonii and A. argen-
teum it was demonstrated that the highest germina-
tion percentage occurred when seeds were 8-10
months old [9, 10, 22]. The same authors consider
that the seeds have lost their viability rather early (af-
ter 20 months). The germinability of the seeds of
both studied species was similar to that observed in
A. bertolonii and A. argenteum [9].
The decrease in germinability could be due to
loss of viability because of decreased energy gener-
ation by the embryo [23]. The values of the parame-
ters used for evaluation of the germination process
(MT, CV, MR, U and Z) also confirmed low germi-
nability and differences in germination process be-
tween the populations of the two Ni-hyperaccumula-
tors. The clearest difference between species and
studied populations was found in the mean germina-
tion time and the related with this parameter mean
germination rates. They were probably related to the
specifics in physiology of the seeds in both species.
These specifics possibly include two groups of fac-
tors - environmental (temperature, water, light, in-
hibitors, etc.) and factors within the seed itself, i.e.
its dormancy which needs future detailed analyses.
The asynchronous germination process spread
through time in A. markgrafii was at the opposite of
quick and synchronous germination in A. murale
populations. The gradual germination process for A.
markgrafii appeared as one of the main differences
between the species and could be considered a type
of relative dormancy as mentioned for some species
from Malvaceae distributed in dry regions of Brazil
[24]. Also, altitudinal differences in the distribution
of both species and the ecological conditions pro-
vided by the environment could be the reason for
these results as mentioned for other plant groups [25,
26]. Similarly to the data for A. argenteum [9], a con-
centration of 1 mM Ni slightly stimulated the germi-
nability in both species while the higher concentra-
tions (2-8 mM Ni) induced a marked inhibition of the
germination process. The inhibition was different
not only between the species but also between their
populations. The seed germination in Ni-hyperaccu-
mulator A. murale was less influenced compared to
Ni-hyperaccumulator A. markgrafii.
A comparison between the root lengths in A.
murale populations showed that the population from
Prrenjas was more tolerant as the mean root length
in all treatments was higher while for A. markgrafii
it was the population from Gjegjan. Seeds root elon-
gation in both species decreased approximately 50%
1350
Fresenius Environmental Bulletin
compared to the controls when a concentration of 1
mM Ni was applied. Similar effects were reported
for Eruca seeds [27]. The results confirm the data of
Vinterhalter and Vilterhalter [28] that tolerance to Ni
is a specific feature to the cells and tissues of certain
species and not only a result of metal translocation.
Even in concentrations below 1 mM a stimulation
effect in root elongation was not found at the oppo-
site of the data for Eruca [26]. The stimulation effect
of Ni was demonstrated in hypocotyl elongation for
all studied populations when the seeds were treated
with Ni concentrations below 1 mM. In all species
and populations the roots were more sensitive to Ni
compared to hypocotyl and their length was lower.
The applied concentrations of 8 mM Ni in our
study reduced significantly both root and hypocotyl
elongation which was evident effect of metal toxicity
in plants. The pattern of growth in root and hypo-
cotyl were different in both species. Seed hypocotyl
growth was less pronounced compared to the root.
Seed root exhibited slowed elongation highly ex-
pressed with the increase of Ni concentrations when
elongation almost stopped. Cell division at the root
tip and cell elongation in the extension zone are con-
sidered two different mechanisms in root growth
both affected by the presence of Ni [7]. The lack of
root hairs, found as most frequent abnormality, could
be considered as one of the reasons for lower nutrient
absorption already mentioned for plants treated with
Ni and smaller size of seedlings of Alyssum species
treated with Ni higher than 1 mM [12]. The physio-
logical regulation of nutrient uptake in serpentine
plants is in relation to Ca and Mg. In serpentine
plants cessation of cell wall extension, disintegration
of cell walls and tissue collapse (necrosis) due to
pectin breakdown, which is most apparent in actively
growing tissues like root meristem, are also consid-
ered typical symptoms of Ca-deficiency [29]. Excess
Ni also affects plant metabolism, inhibits photosyn-
thesis and transpiration, and causes ultrastructural
modifications [12, 30].
CONCLUSIONS
In this study for the first time are demonstrated
differences in the effects of nickel on seed germina-
tion, root and hypocotyl elongation of the facultative
nickel hyperacumulator A. murale and the obligate
nickel hyperaccumulator A. markgrafii, both with
potential in green technologies such as phytoremedi-
ation and phytomining. The laboratory tests for seed
germination showed that in spite of their hyperaccu-
mulation abilities toxic Ni levels inhibited the seed
germination process. The process of elongation was
almost completely inhibited when 8 mM Ni was ap-
plied. Even when treated with low concentrations of
Ni some abnormalities appeared at an early stage of
seed germination. They disturb the normal growth of
© by PSP Volume 27 – No. 3/2018 pages 1345-1352
the seedlings and finally can reduce yields from
these plants.
ACKNOWLEDGEMENTS
This study was realized within the ERAS-
MUS+ contract for the academic year 2015/2016 be-
tween the Faculty of Biology at Sofia University,
Bulgaria and the Agricultural University of Tirana,
Albania.
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