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\begin{document}

\title{Manganese accumulation and foliar distribution in the Australian
hyperaccumulators Gossia bidwillii and Gossia acmenoides}



\author[1]{Farida Abubakari}%
\author[1]{Philip Nti Nkrumah}%
\author[2]{Dennis R Fernando}%
\author[3]{Peter D Erskine}%
\author[1]{Guillaume  Echevarria}%
\author[1]{Antony Van der Ent}%
\affil[1]{The University of Queensland}%
\affil[2]{La Trobe University}%
\affil[3]{Universtiy of Queensland}%


\vspace{-1em}



  
  \date{May 26, 2024}


\begingroup
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\selectlanguage{english}
\begin{abstract}
The known manganese (Mn) hyperaccumulator Gossia bidwillii, and G.
acmenoides unknown to hyperaccumulate Mn are tree species native to
subtropical eastern Australia, where they co-occur on Mn-rich soils.
Here, we investigate Mn accumulation and distribution in G. acmenoides
collected from its natural habitat, and propagated G. bidwillii plants
in a Mn dosing trial. Gossia bidwillii were subjected to different
levels of Mn (250 \selectlanguage{greek}µ\selectlanguage{english}g g-1, 500 \selectlanguage{greek}µ\selectlanguage{english}g g-1, 1000 \selectlanguage{greek}µ\selectlanguage{english}g g-1) treatments whereas G.
acmenoides were sampled from Mn rich soil. We used laboratory-based
X-ray Florescence Microscopy (XFM) to elucidate in situ distribution
patterns of Mn and other elements in hydrated G. acmenoides and G.
bidwillii tissues. Data from G. acmenoides revealed that contrary to
existing knowledge, it can be strongly Mn-hyperaccumulating, with foliar
Mn concentrations of 39 000 \selectlanguage{greek}µ\selectlanguage{english}g g-1 and 24 000 \selectlanguage{greek}µ\selectlanguage{english}g g-1 in old and young
leaves, respectively. In the Mn dosing trial, G. bidwillii accumulated
24 400 \selectlanguage{greek}µ\selectlanguage{english}g g-1 in old leaves and 17 100 \selectlanguage{greek}µ\selectlanguage{english}g g-1 in young leaves at the
highest treatment level. The XFM data revealed clear interspecies
differences in foliar Mn distribution patterns, with wild G. acmenoides
leaves uniformly enriched throughout the laminae and petioles of both
young and old leaves; while in G. bidwillii, the foliar Mn distribution
was primarily concentrated at the apex and lamina. The approach employed
of combining data from the field and controlled experiments was
especially effective for comparing Mn accumulation in these two species
and gaining added insight into the phenomenon of Mn hyperaccumulation.%
\end{abstract}%



\sloppy


\textbf{Manganese accumulation and foliar distribution in the Australian
hyperaccumulators \emph{Gossia} \emph{bidwillii} and \emph{Gossia
acmenoides}}

Farida Abubakari\textsuperscript{1}, Philip Nti
Nkrumah\textsuperscript{1}, Denise R. Fernando\textsuperscript{2}, Peter
D. Erskine\textsuperscript{1},

Guillaume Echevarria\textsuperscript{1,3}, Antony van der
Ent\textsuperscript{1,3*}

\textsuperscript{1}Centre for Mined Land Rehabilitation, Sustainable
Minerals Institute,

The University of Queensland, Australia.

\textsuperscript{2}Department of Ecology, Environment and Evolution, La
Trobe University, Australia.

\textsuperscript{3}Universit\selectlanguage{ngerman}é de Lorraine-INRAE, Laboratoire Sols et
Environnement, 54000 Nancy, France.

* Corresponding author~: a.vanderent@uq.edu.au

\section*{Abstract}

{\label{abstract}}

The known manganese (Mn) hyperaccumulator \emph{Gossia bidwillii} ,
and\emph{G. acmenoides} unknown to hyperaccumulate Mn are tree species
native to subtropical eastern Australia, where they co-occur on Mn-rich
soils. Here, we investigate Mn accumulation and distribution in \emph{G.
acmenoides} collected from its natural habitat, and propagated \emph{G.
bidwillii} plants in a Mn dosing trial. \emph{Gossia bidwillii} were
subjected to different levels of Mn (250 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1}, 500
\selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1}, 1000 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1}) treatments
whereas \emph{G. acmenoides} were sampled from Mn rich soil. We used
laboratory-based X-ray Florescence Microscopy (XFM) to elucidate\emph{in
situ} distribution patterns of Mn and other elements in hydrated\emph{G.
acmenoides} and \emph{G. bidwillii} tissues. Data from \emph{G.
acmenoides} revealed that contrary to existing knowledge, it can be
strongly Mn-hyperaccumulating, with foliar Mn concentrations of 39 000
\selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} and 24 000 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} in old and
young leaves, respectively. In the Mn dosing trial, \emph{G.
bidwillii}accumulated 24 400 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} in old leaves and
17 100 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} in young leaves at the highest treatment
level. The XFM data revealed clear interspecies differences in foliar Mn
distribution patterns, with wild \emph{G} . \emph{acmenoides} leaves
uniformly enriched throughout the laminae and petioles of both young and
old leaves; while in \emph{G. bidwillii} , the foliar Mn distribution
was primarily concentrated at the apex and lamina. The approach employed
of combining data from the field and controlled experiments was
especially effective for comparing Mn accumulation in these two species
and gaining added insight into the phenomenon of Mn hyperaccumulation.

\textbf{KEYWORDS}

elemental distribution, hyperaccumulator, manganese, \emph{Gossia
bidwillii, Gossia acmenoides}

\textbf{INTRODUCTION}

Hyperaccumulators are unique plants able to concentrate extraordinarily
high concentrations of specific trace elements in their foliage and
other aerial parts (Baker \& Brooks, 1989; Reeves, 2003; van der
Ent\emph{et al.} , 2013). They achieve such extreme levels of
accumulation\emph{via} enhanced uptake and translocation mechanisms that
are yet to be fully understood (Baker, 1981; Baker, 1987). Manganese
(Mn) hyperaccumulation is recognised at the notional threshold
concentration of 10 000 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Mn in dry weight shoot
tissue (van der Ent et al., 2013). The hyperaccumulation of Mn is a rare
trait documented primarily within the genera \emph{Alyxia} ,
\emph{Denhamia (Maytenus)} , \emph{Gossia,} \emph{Grevillea,}
\emph{Macadamia} and\emph{Virotia} distributed over eastern Australia
and New Caledonia (Losfeld et al., 2015, Fernando et al., 2008, Jaffr\selectlanguage{ngerman}é,
1977, Jaffré, 1980, Fernando et al., 2009b), Malaysia (Nkrumah et al.,
2018) and recently from Papua New Guinea (Do et al., 2019).

There are 20 Australian \emph{Gossia} species with a wide latitudinal
distribution, ranging from northern New South Wales
(32\textsuperscript{0}S) to the northern tip of the Cape York Peninsula
(10\textsuperscript{0}S) in Queensland (Snow et al., 2003). \emph{Gossia
bidwillii} and \emph{G. acmenoides} (Myrtaceae) have a very smooth bark
which is irregularly covered with relatively large, coloured patches.
These species, as well as \emph{G. lucida} and \emph{G. grayi,} are
called ``python bark'' \emph{Gossia's} due to the resemblance of their
bark to the skin colouring of the python snake (Snow et al.,
2003).\emph{Gossia bidwillii} , is the only Australian \emph{Gossia} to
thrive on ultramafic soils (McLay et al., 2019), and also the first Mn
hyperaccumulator described in Australia (Bidwell \emph{et al.} , 2002).
That discovery instigated subsequent research on Mn hyperaccumulation in
several other Australian \emph{Gossia} species (Fernando \emph{et al.} ,
2007; Fernando \emph{et al.} , 2008a; Fernando \emph{et al.} , 2009b;
Fernando \emph{et al.} , 2013, McLay 2018). All these published studies
have been based on freshly collected field samples or preserved material
obtained from herbaria. There have been certain consistent observations
throughout, for example, the Mn hyperaccumulative trait in \emph{Gossia
bidwillii} , and the Mn non-hyperaccumulation by \emph{G. acmenoides,}
as captured in a recent phylogenetic study of \emph{Gossia} (McLay et al
2018). While these two species are known to be sympatric on Mn-rich
soils, they appear taxonomically partitioned into separate clades
(McClay et al (2018).

Studies on fresh field material have reported foliar Mn concentrations
of 19 200 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} in \emph{G. bidwillii} (Bidwell et
al., 2002). Recent growth experiment on \emph{G. fragrantissima} has
shown it can take up to 545 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Co, 17 400 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} Mn and up to 13 000 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Zn
(Abubakari et al. 2021a) whereas freshly collected field samples
of\emph{G. grayi} and \emph{G. shepherdii} were observed to contain up
to 13 700 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} and 11 000 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1}foliar Mn respectively (Fernando et al., 2018).
\emph{In vivo}cryo-scanning electron microscopy (SEM)/energy dispersive
X-ray analysis (EDS) showed Mn localization in\emph{G. bidwillii} to be
different from other hyperaccumulating species. Foliage hyperaccumulated
metals are usually known to accumulate in non-photosynthetic tissues
such as the epidermal cells and associated dermal structures including
trichomes and leaf hairs (V\selectlanguage{ngerman}ázquez et al., 1992, Küpper et al., 2000,
Küpper et al., 2001, Krämer et al., 1997, Mesjasz-Przybylowicz et al.,
2001, Bhatia et al., 2003, Bidwell et al., 2004, Broadhurst et al.,
2004), whereas in \emph{G. bidwillii} Mn was found to be primarily
localised in photosynthetic cells (Fernando et al., 2006b, Fernando et
al., 2006a, Fernando et al., 2007). Laboratory and synchrotron X-ray
Florescence Microscopy (XFM) have revealed marginal accumulation of Co,
Mn and Zn in leaves, with localization of Co, Mn and Zn in epidermal
cells of \emph{G. fragrantissima} (Abubakari et al. 2021a).

To date, no attempt has been made to examine the effects of Mn dosing
treatments on \emph{G} . \emph{bidwillii} under controlled experimental
conditions, mainly due to the relatively slow growth rate of woody
species such as this. Neither have there been any studies to examine Mn
accumulation of \emph{G. bidwillii} to Mn accumulation by a closely
related species. \emph{Gossia acmenoides} was selected here for a field
investigation into Mn accumulation in its natural habitat because it is
commonly sympatric to \emph{G. bidwillii} . This study aims to: i)
measure the response of propagated \emph{G. bidwillii} plants to Mn
treatment under controlled conditions, and ii) assess Mn uptake and
accumulation in the aforementioned \emph{G. bidwillii} plants with that
of \emph{Gossia acmenoides} on naturally Mn enriched soils\emph{.} By
employing laboratory XFM to determine \emph{in situ} distributions of Mn
and other elements in \emph{G. bidwillii} and \emph{G.acmenoides,} this
study also investigates Mn distribution patterns in the leaf tissues of
these two species, as well as test within-species age-related
distributional differences, \emph{i.e} ., between their young and old
leaves.

\textbf{MATERIALS AND METHODS}

\textbf{Habitat and ecology of the studied species} \emph{-- Gossia
acmenoides} and \emph{G. bidwillii} are small to medium trees up to
\textasciitilde{}18 m tall, which occur in drier scrub patches and
rainforests. \emph{Gossia acmenoides} has a brown/green bark, shedding
in patches; terminal buds silky. Leaves are oval to elliptical quickly
tapering to a pointed or rounded tip, simple, opposite, blade moderately
glossy above, translucent, venation distinct, old dots visible with the
naked eye (Snow et al., 2003). Unlike \emph{G. bidwillii} , the leaves
of\emph{G. acmenoides} are not sticky when crushed. It is a hardy plant
with dense and glossy foliage. On the other hand, \emph{G}
.\emph{bidwillii} have almost round, very smooth, shiny and a cinnamon
smell when crushed (Fig. 1).

\textbf{Plant dosing trial of \emph{G. bidwillii --}} \emph{Gossia
bidwillii} plants, approximately 20 cm in height, were obtained from
Coastal Dry Tropics Landcare (Pallarenda Road, Townsville, Queensland)
and cultivated in a temperature and humidity-controlled glasshouse.
Plants were kept at 20 ºC and 80\% Relative Humidity (RH), with and
13:00 hours of PAR light (1600 \selectlanguage{greek}μ\selectlanguage{english}M photons s\textsuperscript{-1}) at the
Central Glasshouse Services at The University of Queensland, Brisbane,
Australia. After three weeks, the plants were transferred into 15 cm
pots containing a ratio of 9:1 mixture of Composted Pine Bark 5--10 mm
and Coco Peat (Bassett Barks Pty Ltd, Queensland, Australia). The media
was mixed with low-level fertilizers and other augments consisting (per
m\textsuperscript{3}) of 1.2 kg Yates Flowtrace, 1 kg iron sulphate
heptahydrate
(FeO\textsubscript{4}SO\textsubscript{4}\selectlanguage{ngerman}·7H\textsubscript{2}O), 0.1 kg
superphosphate
(Ca(H\textsubscript{2}PO\textsubscript{4})\textsubscript{2}), 1.5 kg
gypsum (CaSO\textsubscript{4}) and 1.5 kg dolomite
(CaMg(CO\textsubscript{3})\textsubscript{2}). The composition of the
Flowtrace was 24 wt\% iron (Fe) as FeSO\textsubscript{4}, 14 wt\% sulfur
(S) as SO\textsubscript{4}, 0.75 wt\% copper (Cu) as
CuSO\textsubscript{4}, 0.5 wt\% manganese (Mn) as MnSO\textsubscript{4},
0.2 wt\% zinc (Zn) as ZnSO\textsubscript{4}, 0.04 wt\% molybdenum (Mo)
as Na\selectlanguage{english}2MoO4, \selectlanguage{english}0.033 wt\% boron (B) as
Na\textsubscript{2}B\textsubscript{4}O\textsubscript{7} and also
contains zeolite, to ensure flowability (Yates Australia, Padstow, NSW,
Australia). Soluble Mn was applied to the plants in a randomised block
design. The applied treatments were the control (\textbf{T1} ), and
soils with final dosed Mn\textsuperscript{2+} concentrations of 200 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} (\textbf{T2} ), 500 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1}(\textbf{T3} ) and 1000 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1}
(\textbf{T4} ) replicated three times yielding a total of 12
experimental groups. Each treatment was administered monthly as aqueous
MnSO\textsubscript{4}.H\textsubscript{2}O solutions for a period of 12
months; a similar volume of water was added to the control each time.
The individual pots were placed on saucers and hand watered daily to
field capacity to prevent loss of treatment solutions.

\textbf{Field sampling of \emph{G. acmenoides}} -- \emph{Gossia
acmenoides} was sampled at a field site within the Amamoor State Forest
in Queensland, Australia (26\selectlanguage{ngerman}°20'41.0''S 152°37'7.0''E). The geology of
this subtropical area is predominantly volcanic rock (andesite)
overlaying variably silicified shale or tuffs containing Mn-rich
(\textasciitilde{}30--50 wt\%) minerals such as bixbyite and pyrolusite.
Krasnozem soils derived from this parent rock contain Mn to levels as
high as 40 wt\% (Isbell, 1994). Old and young \emph{G. acmenoides}leaves
(10--20 each) were harvested for total mineral nutrient analysis, while
small branchlets with old and young leaves were detached and stored
fresh for XRF analyses. Soil samples were collected from beneath the
trees (\textless{}10 cm depth) at three different points free of surface
litter.

\textbf{Chemical analysis of soil and plant samples} -- After
harvesting\emph{G. bidwillii} , soils were extracted in each pot and
emptied into respective plastic bags. Soils on which \emph{G.
acmenoides} was growing were also collected as described above. All
soils were oven dried at 60º C and later sieved using the 2mm sieve.
Soil pH was obtained in a 1 to 2.5 soil to water mixture after 2 hr
shaking. Exchangeable trace elements were extracted in 0.1 M
Sr(NO\textsubscript{3})\textsubscript{2} at a soil:solution ratio of 1:4
(10 gram soil with 40 mL solution) and 2 hr shaking time was adapted
from Kukier and Chaney (2001). As a means of estimating potentially
phytoavailable trace elements, the DTPA-extractant was used according to
Dai \emph{et al.} (2004) which was adapted from the original method by
Lindsay and Norvell (1978), with the following modifications: excluding
TEA, adjusted at pH 5.3, 5 g soil with 25 mL extractant, and extraction
time of one hr.

Plant material samples were oven dried at 60°C for three days and then
weighed, ground to fine powder and (300 mg) digested using 4 mL
HNO\textsubscript{3} (70\%) in a microwave oven (Milestone Start D) for
a 45-minute programme. Digests were then diluted to 45 mL with ultrapure
water (Millipore 18.2 M\selectlanguage{greek}Ω\selectlanguage{ngerman}·cm at 25°C) for analysis with Inductively
coupled plasma atomic emission spectroscopy (ICP-AES) using a Thermo
Scientific iCAP 7400 instrument for macro-elements (Al, Na, Mg, K, P,
Ca) and trace-elements (Fe, Ni, Mn, Co, Zn) in radial and axial modes,
depending on the element and expected analyte concentration. In-line
internal standardization using yttrium was used to compensate for
matrix-based interferences. Quality controls included matrix blanks,
certified reference material (Sigma-Aldrich Periodic Table mix 1 for ICP
TraceCERT®, 33 elements, 10 mg L\textsuperscript{-1} in
HNO\textsubscript{3}) and Standard Reference Material (NIST Apple 1515
digested with HNO\textsubscript{3}).

\selectlanguage{greek}\textbf{Laboratory µXRFelemental mapping}\selectlanguage{english} -- Live samples (a whole
branch) from \emph{Gossia bidwillii}originating from the Mn1000
treatment and \emph{G. acmenoides} collected from Amamoor, Queensland
were used for the microXRF scanning. The UQ microXRF facility contains a
modified IXRF ATLAS X system, mounting two 50W X-ray sources fitted with
polycapillary focussing optics: XOS microfocus Mo-target tube producing
17.4 keV X-rays (flux of 2.2 \selectlanguage{ngerman}× 10\textsuperscript{8} ph
s\textsuperscript{-1}) focussing to 25 \selectlanguage{greek}μ\selectlanguage{english}m and a Rh-target tube producing
20.2 keV (flux of 1.0 \selectlanguage{ngerman}× 10\textsuperscript{7}ph s\textsuperscript{-1})
focussing to 5 \selectlanguage{greek}μ\selectlanguage{english}m. The system is fitted with two silicon drift detectors
of 150 mm\textsuperscript{2}. Typical energy resolution is
\textless{}145 eV with a maximum input count rates of 2 M counts per
second. The motion stage can address areas up to 300 \selectlanguage{ngerman}× 300 mm.
Measurements were conducted at atmospheric temperature
(\textasciitilde{}20°C), using the Mo 25 \selectlanguage{greek}μ\selectlanguage{english}m X-ray source at a 40 kV,
1000 uA, with a rise time of 0.25 \selectlanguage{greek}µ\selectlanguage{english}s and a per-pixel dwell of 100 ms.
The hydrated foliar samples were mounted between two sheets of 4 \selectlanguage{greek}μ\selectlanguage{english}m
Ultralene thin film in a tight sandwich to limit evaporation and
analysed within 10 minutes after excision. The mounted samples between
Ultralene thin film were stretched over a Perspex frame magnetically
attached to the x-y motion stage at atmospheric temperature
(\textasciitilde{}20\selectlanguage{ngerman}°C). The possibility of radiation-induced damage in
\selectlanguage{greek}μ-\selectlanguage{english}XRF analysis (especially in fresh hydrated samples) is an important
consideration, but such damage was not observed because the source
produced a flux of 2.2 \selectlanguage{ngerman}× 10\textsuperscript{8}photons
s\textsuperscript{-1} in a 25 \selectlanguage{greek}μ\selectlanguage{english}m beam spot, at a maximum dwell of 100 ms
this results in a deposited radiation dose of just 6.6 Gy.

\textbf{Data processing and statistical analysis} -- The XRF spectra on
the UQ microXRF facility were acquired in mapping mode using the
instrument control package, Iridium (IXRF systems) from the sum of
counts at the position of the principal peak for each element. These
were each exported into ImageJ as greyscale 8-bit TIFF files, internally
normalised such that each image covered the full dynamic range and
displayed using ImageJ's ``Fire'' lookup table.

The concentrations of Mn presented as boxplots were performed using R
version 3.6.1 (2019-07-05). Concentrations of elements presented in
Tables as mean \selectlanguage{ngerman}± standard error were conducted using One-Way ANOVA and
means compared with Tukey's honestly significant difference (HSD) Post
Hoc Test in the IBM SPSS Statistics 27 software package (IBM, New York,
USA). Values with different small letters are significantly different
(\emph{p} \textless{}0.05).

\textbf{RESULTS}

\textbf{Elemental concentrations in \emph{G. acmenoides} and \emph{G.
bidwillii}} -- All plant tissue elemental concentrations presented in
this study are based on dry weight. The bulk elemental concentrations in
young leaves, old leaves and twigs of the wild \emph{G.acmenoides} and
the dosed \emph{G. bidwillii} are shown in Table 1 and 2 respectively,
and that of the concentration of Mn in young, old leaves and twigs of
\emph{G. bidwillii} in Figure 2. The concentrations of Mn in the wild
\emph{G. acmenoides} were remarkable, with significantly (\emph{p}
\textless{}0.05) high mean value of 39 000 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE
\selectlanguage{ngerman}± 3540) in old leaves, compared to 24 000 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE \selectlanguage{ngerman}±
410) in young leaves and 5840 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 2820) in
twigs (Table 1). Similarly, the concentrations of Ca and Na in \emph{G.
acmenoides} tend to be high in old leaves (9250 ± 785 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} Ca, 710 \selectlanguage{ngerman}± 70.0 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Na) than
in young leaves (5590 \selectlanguage{ngerman}± 145 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Ca, 660 \selectlanguage{ngerman}± 190 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} Na) and twigs (4700 \selectlanguage{ngerman}± 400 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1}
Ca, 170 \selectlanguage{ngerman}± 35.0 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Na) (Table 1). The
concentrations of Fe and Mg in old leaves of \emph{G.} \emph{acmenoides}
were also high with concentrations of 75.0 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Fe
(SE \selectlanguage{ngerman}± 17.0) and 1290 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Mg (SE \selectlanguage{ngerman}± 108) respectively
compared to 840 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Mg (SE \selectlanguage{ngerman}± 38.0) and 45 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} Fe (SE \selectlanguage{ngerman}± 0.70) in its young leaves. However, no
significant difference was observed between Mg in old leaves and twigs
of \emph{G.acmenoides} (Table 1). Values of Al tend to be also high in
old leaves (200 ± 38.0 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Al) than young leaves
(53.0 \selectlanguage{ngerman}± 0.85 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Al) and twigs (6.0 \selectlanguage{ngerman}± 10.5 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} Al). Nonetheless, the concentration of K was high
in young leaves (9380 \selectlanguage{ngerman}± 1040 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} K) than old leaves
(3660 \selectlanguage{ngerman}± 160 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} K) and twigs (2710 \selectlanguage{ngerman}± 570 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} K). No significant difference
(\emph{p}\textgreater{}0.05) was observed for Ni, P and Zn in young, old
leaves and twigs of \emph{G. acmenoides} (Table1).

In the old leaves of \emph{G. bidwillii} , the mean Mn concentrations at
the \textbf{T4} , \textbf{T3} and \textbf{T2} treatment levels were 24
400 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 10 700), 21 800 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 8850) and 14 100 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1}(SE
\selectlanguage{ngerman}± 12 600), respectively. These are more than 2-fold the Mn
hyperaccumulation threshold at the \textbf{T4} and \textbf{T3} treatment
levels and greater than 1-fold the Mn hyperaccumulation threshold at
the\textbf{T2} treatment level (Fig. 2 and Table 2). The old leaves
of\emph{G. bidwillii} at the control (\textbf{T1} ) also contained up to
10 900 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Mn (mean 5490 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1}).
In the young leaves, however, Mn concentration at the \textbf{T4}
and\textbf{T3} treatment levels were 17 100 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE
\selectlanguage{ngerman}± 8440) and 12 600 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 11 000) respectively
which are more than 1-fold the Mn hyperaccumulation threshold (Fig. 2
and Table 2). On the other hand, the \textbf{T2} treatment level
contained 8300 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 4270) which is almost
around double the \textbf{T1} value (4600 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1}) in
young leaves (Fig. 2 and Table 2). In the twigs, the highest treatment
level (\textbf{T4} ) contained 17 400 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE \selectlanguage{ngerman}±
6690) Mn which is more than 1.5-fold the Mn hyperaccumulation threshold
than that observed at the \textbf{T3} (5000 ± 1150 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1}), \textbf{T2} (4300 \selectlanguage{ngerman}± 1220 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1}) and the \textbf{T1} (2900 \selectlanguage{ngerman}± 1300 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1}) treatment levels (Fig. 2 and Table 2).

Unlike \emph{G. acmenoides} , the concentration of Ca in the
dosed\emph{G. bidwillii} was high in the young leaves with
concentrations of 9400 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 2280) at
\textbf{T1} , 5700 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 1820) at \textbf{T2} ,
12 400 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 2790) at \textbf{T3} and 11 200 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 4000) at \textbf{T4} treatment level
compared to that in the old leaves (Table 2). The twigs on the other
hand, contained high amounts of Ca at the \textbf{T1} , \textbf{T2}
and\textbf{T4} treatment levels than in young and old leaves with mean
values of 17 100 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 1580), 11 100 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 2970) and 11 700 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1}(SE
\selectlanguage{ngerman}± 3780) respectively (Table 2) except at the \textbf{T3} treatment level
where the concentration of Ca was high in young leaves than in old
leaves and twigs. The concentration of Al in old leaves of \emph{G.
bidwillii} tend to be high at the \textbf{T3} and \textbf{T4} treatment
levels with concentrations of 9000 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 5400)
and 10 700 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (SE \selectlanguage{ngerman}± 5370) respectively compared to
that in the young leaves and twigs (Table 2). The concentration of K at
the \textbf{T1} (22100 ± 2960 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} K) treatment
level in young leaves was significantly (\emph{p} \textless{}0.05)
higher than those observed at the other treatment levels (\textbf{T2}
,\textbf{T3} , \textbf{T4} ) in old leaves and twigs. However, Mg tend
to be high at the \textbf{T1} (4790 \selectlanguage{ngerman}± 1900 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} Mg)
treatment level in old leaves compared to those observed at the other
treatment levels (\textbf{T2} , \textbf{T3} , \textbf{T4} ) in young
leaves and twigs (Table 2). Whereas no significant difference was
observed between the concentration of Na at the \textbf{T1} and
\textbf{T2}treatment levels of young and old leaves, the concentration
of Na at the\textbf{T4} treatment level in twigs (1100 \selectlanguage{ngerman}± 840 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} Na) was significantly (\emph{p} \textless{}0.05)
higher compared to that in young leaves (440 \selectlanguage{ngerman}± 260 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} Na) and old leaves (630 \selectlanguage{ngerman}± 140 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1} Na) (Table 2). The values for Fe and Zn were low
at all treatment levels compared to other elements in young, old leaves
and twigs (Table 2). Nonetheless, the concentration of P was below the
limit of detection (\textless{}6.00 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1}) in young,
old leaves and twigs at all treatment levels, and this could probably be
due to the low P demand in the experimental \emph{G. bidwillii} (Table
2).

\textbf{Soil chemical properties} -- The soil pH of the wild \emph{G.
acmenoides} was low (pH = 4.20) compared to that of \emph{G.
bidwillii}pot soils (pH 5.27--5.95) (Table 3). Extractable Mn values for
DTPA and Sr(NO\textsubscript{3})\textsubscript{2} increased with the
dose levels, as expected. Variability in Mn values for DTPA and
Sr(NO\textsubscript{3})\textsubscript{2} can be explained by the strong
binding of Mn\textsuperscript{2+} to the organic matter of the potting
mix thereby making it less available.

\selectlanguage{greek}\textbf{µXRF elemental mapping}\selectlanguage{english} \textbf{of wild
\emph{G}.\emph{acmenoides} and \emph{G. bidwillii}} -- Manganese in the
wild\emph{G} . \emph{acmenoides} collected from the Mn-enriched
substrate in Amamoor has strong enrichment throughout the leaf blade and
petiole of young and old leaves, while veins and midrib have relatively
lower concentrations (Figs. 3 and 4). Calcium and K are strongly
enriched in the veins and midrib, and with high concentrations of Ca in
the stem and petiole and low K in the leaf margins of young and old
leaves (Fig. 3). In \emph{G. bidwillii} , the distribution of Mn is high
at the apex and lamina but low in the midrib, margin and veins (Fig. 4).
However, K tends to be concentrated at the tip/margin, midrib, and
petiole and veins whereas Ca is high in the midrib, margin and veins but
low in the leaf lamina (Fig. 4).

\textbf{DISCUSSION}

This study details the first-ever growth experiment on the Mn
hyperaccumulating tree, \emph{G. bidwillii} , exposed here to a range of
Mn treatments and reports for new discovery of the Mn hyperaccumulative
trait in wild \emph{G. acmenoides} , on the highly acidic Mn-rich soils
is intriguing given past analyses of samples sourced from a range of
other eastern Australian locations had not detected Mn hyperaccumulation
in this species (Fernando et al. 2009b). Foliar Mn concentrations in
experimental \emph{G. bidwillii} to as high as \textasciitilde{}3-fold
Mn hyperaccumulation threshold of 10~000 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (van
der Ent et al. 2013) aligns with field data from earlier studies of this
species (Bidwell et al. 2002; Fernando et al. 2006b). A weak
relationship between substrate Mn supply and foliar Mn concentrations,
most notably, high Mn uptake by control plants is also consistent with
past field observations (Bidwell et al., 2002, Fernando et al., 2006b,
Fernando et al., 2007). The ability of \emph{Gossia bidwillii} as shown
experimentally here and as previously noted in the field to vastly
over-accumulate Mn even in very low soil-supply is a common
characteristic hyperaccumulation, \emph{i.e.,} the ability to scavenge
from host substrates (Baker 1981; (Fernando et al., 2007).

Contrasting findings of previous field studies that have not
found\emph{G. acmenoides} to hyperaccumulate Mn, and this present field
study describing strong Mn hyperaccumulation by this species may be
attributable to genetic differences, as also observed in \emph{Denhamia
founieri} from New Caledonia by Fernando et al. (2008). The present
findings on \emph{G. acmenoides} suggests heterogeneity across the
species, warranting further investigation of the genetic basis of Mn
hyperaccumulation in \emph{Gossia,} for example, as interrogated by
Pollard et al (2002) for ubiquitous metal hyperaccumulating herbs of the
Northern Hemisphere. Variability of Mn hyperaccumulation trait
in\emph{G. bidwillii} has previously been reported, however there is
greater consistency across its broad natural range in comparison to the
emerging picture of \emph{G. acmenoides} (Fernando et al., 2007).
Heterogeneity of metal accumulation has also been described in several
species of hyperaccumulators of metals other than Mn (Pollard et al.,
2002, Baker et al., 1994, Macnair, 2002). The extreme acidity of the
host soil (pH 4.20, Table 3) from which \emph{G. acmenoides} was sampled
for this study warrants consideration in the context of Mn availability
at the root-soil interface. It is plausible that the apparently high
variation in Mn accumulation by \emph{G. acmenoides} reflects specific
rhizosphere effects such as acidification and/or microbial associations
unique to a particular site location that renders soil-Mn highly
bioavailable. These findings yet again highlight the gaps in knowledge
around metal hyperaccumulation, with woody species such as
\emph{Gossia}spp\emph{.} poorly understood. At locations where \emph{G.
acmenoides}has previously been found to accumulate\textbackslash{}souts
low levels of Mn, there may also be ion competition among similar
divalent cations such as Ca\textsuperscript{2+}, Mg\textsuperscript{2+}
and Mn\textsuperscript{2+} in addition to less acidic soil conditions
(Fernando et al., 2013, McLay et al., 2019, Bidwell et al., 2002). It is
notable that high concentrations of Ca and Mg relative to Mn were
reported in leaves of \emph{G. acmenoides} (Bidwell et al. (2002).

In \emph{G. bidwillii} , the behaviour of Mn in the oldest leaves
resembled that of Ca, Mg and Na which remained high at all treatment
levels but contrasted with that of K and P which decreased after
maturity. However, in \emph{G. acmenoides} , K was high in young leaves
than in old leaves, whereas Ca, Mg, Na and Mn were higher in old leaves.
The high concentrations of Mn, Ca, Mg and Na in old leaves of \emph{G.
bidwillii} and \emph{G. acmenoides} could be attributed to phloem
immobility of the aforementioned elements (Marschner, 2002, Graham et
al. 1988) and vice versa for K in \emph{G} . \emph{acmenoides}
and\emph{G. bidwillii} . Moreover, this behaviour could be due to the
similarities in divalent cations of Mn, Ca and Mg. Similar observation
of high Mn in old leaves has been reported in \emph{G.
bidwillii}(Bidwell et al., 2002) and in other Mn hyperaccumulators
including\emph{Phytolacca americana} (Xu et al., 2006), \emph{Macadamia
integrifolia} (Fernando et al., 2009a), \emph{G.
fragrantissima}(Abubakari et al. 2021) and in crop plants (Millikan,
1951). In contrast, \emph{G. grayi} and \emph{G. shepherdii} were
reported to accumulate higher Mn concentrations in young leaves than in
older leaves (Fernando et al., 2018). A previous report by Bidwell et
al. (2002) of decreased Ca and Mg with an increase in Mn concentration
in old leaves of \emph{G. bidwillii} contradicts the findings of this
study. The nutritional dynamics of plants as unusual as metal
hyperaccumulators are yet to be fully understood, and clearly cannot be
assumed to align with broader understanding of plant nutrition largely
drawn from crop models (Marschner 2002).

The phytoavailability of Al and Mn are known to occur in soils of low pH
(\textless{}5), however the pH of soils on which \emph{G. bidwillii} was
cultivated in this present study shows that the solubility of Al was low
(\textgreater{}5), yet \emph{G. bidwillii} was able to take up high
amounts of Al in old leaves which qualifies it as an Al hyperaccumulator
with concentration 6-fold higher than the Al hyperaccumulation threshold
set at 3000 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1} (Jansen et al., 2003, Jansen et
al., 2001). Exceptionally high Al concentrations in old \emph{G.
bidwillii} leaves, even under Mn treatment, could be facilitated by
organic anions involved in Mn transport (Bidwell et al., 2002). This
suggests that \emph{G.} \emph{bidwillii} may be able to take up Al via
an anion channel, a mechanism that appears to be a peculiar trait among
Al tolerant species (Zhang et al., 2001, Ryan et al., 1997, Pi\selectlanguage{ngerman}ñeros and
Kochian, 2001, Kollmeier et al., 2001). It should be cautioned here that
while these observations of Al over accumulation by \emph{G.
bidwillii}have been made under experimental conditions, it has not been
observed in the field, even on lateritic soils rich in Al (Fernando,
Bidwell etc). The notable limit in uptake of Al in \emph{G. acmenoides}
even though its soil pH was suitably low for Al mobilisation for plant
uptake, was most likely due to the very low soil Al concentrations.
Furthermore, it has also been suggested that Al inhibits uptake of Ca
and Mg in non- Al accumulators (Kochian et al., 2005, Ryan and Kochian,
1993, Rengel and Zhang, 2003), and this was found in old leaves
of\emph{G. acmenoides} in this present study. Species within the
Myrtaceae family have been listed to contain Al hyperaccumulators
(Jansen et al., 2003, Jansen et al., 2001) and previous studies by
Fernando et al. (2009b) have shown that other \emph{Gossia} spp.\emph{,}
including\emph{G. hillii} , \emph{G. inophloia} , \emph{G. lewensis} and
\emph{G. macilwraithensis,} can be Al hyperaccumulators after they were
exposed to Al treatments.

The distribution patterns of Mn in leaves in wild \emph{G.
acmenoides}and Mn-dosed \emph{G. bidwillii} have been shown by XFM to be
distinctly different (Figs. 3 and 4). Strong Mn distribution throughout
young and old leaves in wild \emph{G. acmenoides} , while highly
concentrated at the apex and lamina of treated \emph{G. bidwillii}
suggest that Mn movement with the transpiration stream rather than
against it. This indicates that, increasing transpiration rate
throughout the leaves of\emph{G. acmenoides} , and towards the apex and
lamina of \emph{G. bidwillii} led to higher Mn accumulation in those
parts of the studied species. Similar observations to that of \emph{G.
bidwillii} have been reported for the Mn hyperaccumulators
\emph{Acanthopanax sciadophylloides} (Memon et al., 1980) and \emph{G.
fragrantissima}(Abubakari et al., 2021a).

This study newly revealed the strong and facultative Mn
hyperaccumulative trait in \emph{G. acmenoides} and confirmed it
in\emph{G. bidwillii} , demonstrating both species to be strong Mn
hyperaccumulators with concentrations in both young and old leaves well
exceeding the Mn hyperaccumulation threshold. Laboratory based-XFM
revealed distinct Mn distribution patterns in the leaves of \emph{G.
acmenoides} and \emph{G. bidwillii} . Further work should be undertaken
using synchrotron X-ray Florescence Microscopy with more precision and
higher resolution to investigate Mn distribution at the cellular and
subcellular levels in order to elaborate hypotheses for its metabolic
pathways to be elucidated with genetic studies.

\textbf{ACKNOWLEDGEMENTS}

Farida Abubakari is the recipient of a UQ Graduate School Scholarship
(UQGSS) from The University of Queensland. We thank Lachlan Casey
(Centre for Microscopy and Microanalysis at the University of
Queensland) for technical support with the \selectlanguage{greek}µ\selectlanguage{english}XRF analysis. The authors
acknowledge the support of Microscopy Australia at the Centre for
Microscopy and Microanalysis at the University of Queensland.

\textbf{FUNDING}

No funding was received for conducting this study.

\textbf{AUTHOR CONTRIBUTIONS}

FA, PNN and AVDE designed and conducted the experiment. FA and AVDE
collected the samples and undertook the chemical analysis of the
samples. FA, PNN and AVDE performed data processing and analysis. All
authors contributed to writing of the manuscript.

\textbf{CONFLICT OF INTEREST}

The authors have no conflicts of interest to declare that are relevant
to the content of this article.

\textbf{DATA AVAILABILITY STATEMENT}

Data can be accessed from the Dryad Digital Repository.

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Vazquez MD, Barcelo J, Poschenrieder Ch, Madico J, Hatton P, Baker AJM,
Cope GH. 1992. Localization of zinc and cadmium in \emph{Thlaspi
caerulescens} (Brassicaceae), a metallophyte that can hyperaccumulate
both metals. \emph{Journal of Plant Physiology,} 140: 350-355.

Xu X, Shi J, Chen Y, Chen X, Wang H, Perera A. 2006. Distribution and
mobility of manganese in the hyperaccumulator plant \emph{Phytolacca
acinosa} Roxb. (Phytolaccaceae). \emph{Plant and Soil,} 285: 323-331.

Zhang W-H, Ryan PR, Tyerman SD. 2001. Malate-permeable channels and
cation channels activated by aluminum in the apical cells of wheat
roots. \emph{Plant Physiology,} 125: 1459-1472.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}

\textbf{Figure 1.} Visual appearance\textbackslash{}souts of
experimental and field plants: \textbf{A} Experimental \emph{Gossia
bidwillii} , \textbf{B} ; field \emph{G. acmenoides} tree trunk, and
\textbf{C} field \emph{G. acmenoides} foliage\emph{.}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image2/image2}

\end{center}
\end{figure}

\textbf{Figure 2.} Manganese concentrations in young leaves, old leaves,
and twigs of experimental

\emph{Gossia bidwillii} plants. Values are in ranges and means of three
replicates. Keys to symbol of

boxplots: open squares are the 25\% to 75\% quartiles; whiskers are the
mean \selectlanguage{ngerman}± standard error and circles are outliers. Values with different
small letters are significantly different (\emph{p} \textless{}0.05).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image3/image3}

\end{center}
\end{figure}

\textbf{Figure 3} . Laboratory \selectlanguage{greek}µ\selectlanguage{english}XRF maps of K, Ca and Mn of a whole
fresh/hydrated \emph{G. acmenoides}branchlets.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image4/image4}

\end{center}
\end{figure}

\textbf{Figure 4.} Laboratory \selectlanguage{greek}µ\selectlanguage{english}XRF maps of K, Ca and Mn of
fresh/hydrated leaves of \emph{G. acmenoides} (top) and \emph{G.
bidwillii} (below).

\textbf{Table 1.} Elemental concentrations in field samples of \emph{G.
acmenoides} young leaves, old leaves, and twigs. Values are average of
three replicates \selectlanguage{ngerman}± standard error (except for young leaves where n =2).
Values with different small letters in the same column are significantly
different (\emph{p} \textless{}0.05).\selectlanguage{english}
\begin{longtable}[]{@{}lllllllllll@{}}
\toprule
\begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\textbf{Species} \selectlanguage{greek}\textbf{Mn levels (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage}\tabularnewline
\midrule
\endhead
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Al}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Ca}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Fe}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{K}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Mg}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Mn}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Na}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Ni}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{P}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Zn}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
Gossia acmenoides\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\emph{In situ} collection\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
53\selectlanguage{ngerman}±0.85\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
5590\selectlanguage{ngerman}±145\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
45.0\selectlanguage{ngerman}±0.70\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
9380\selectlanguage{ngerman}±1040\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
840\selectlanguage{ngerman}±38.0\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
24 000\selectlanguage{ngerman}±410\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
660\selectlanguage{ngerman}±190\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
10\selectlanguage{ngerman}±1.74\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
445\selectlanguage{ngerman}±83.0\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
8\selectlanguage{ngerman}±1.40\textsuperscript{a}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\emph{In situ} collection\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
200\selectlanguage{ngerman}±38.0\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
9250\selectlanguage{ngerman}±785\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
75.0\selectlanguage{ngerman}±17.0\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
3660\selectlanguage{ngerman}±160\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
1290\selectlanguage{ngerman}±108\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
39000\selectlanguage{ngerman}±3540\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
710\selectlanguage{ngerman}±70.0\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
25\selectlanguage{ngerman}±3.10\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
410\selectlanguage{ngerman}±63.0\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
15\selectlanguage{ngerman}±7.07\textsuperscript{a}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\emph{In situ} collection\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
6.0\selectlanguage{ngerman}±10.5\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
4700\selectlanguage{ngerman}±400\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
17.0\selectlanguage{ngerman}±6.00\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
2710\selectlanguage{ngerman}±570\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
1130\selectlanguage{ngerman}±725\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
5840\selectlanguage{ngerman}±2820\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
170\selectlanguage{ngerman}±35.0\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
90\selectlanguage{ngerman}±85.5\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
440\selectlanguage{ngerman}±60.0\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
20\selectlanguage{ngerman}±9.10\textsuperscript{a}\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}\selectlanguage{ngerman}

\textbf{Table 2.} Elemental concentrations in young leaves, old leaves
and twigs of \emph{G. bidwillii} after exposure to different levels of
Mn treatments. Values are average of three replicates ± standard error.
Values with different small letters in the same column are significantly
different (\emph{p} \textless{}0.05).\selectlanguage{english}
\begin{longtable}[]{@{}lllllllllll@{}}
\toprule
\begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\textbf{Species} \selectlanguage{greek}\textbf{Mn levels (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage} & \begin{minipage}[b]{0.09\columnwidth}\raggedright\strut
\selectlanguage{greek}\textbf{Elemental concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\strut
\end{minipage}\tabularnewline
\midrule
\endhead
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Al}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Ca}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Fe}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{K}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Mg}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Mn}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Na}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Ni}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{P}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Zn}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
Gossia bidwillii\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Young leaves}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
Control\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
20\selectlanguage{ngerman}±12.0\textsuperscript{h}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
9400\selectlanguage{ngerman}±2280\textsuperscript{e}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
40\selectlanguage{ngerman}±18.0\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
22 100\selectlanguage{ngerman}±2960\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
3800\selectlanguage{ngerman}±540\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
4640\selectlanguage{ngerman}±1725\textsuperscript{h}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
1400\selectlanguage{ngerman}±500\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
225\selectlanguage{ngerman}±140\textsuperscript{b}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
200\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
14 200\selectlanguage{ngerman}±7640\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
5700\selectlanguage{ngerman}±1820\textsuperscript{f}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
30\selectlanguage{ngerman}±5.8\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
12 800\selectlanguage{ngerman}±2830\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
3400\selectlanguage{ngerman}±400\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
8300\selectlanguage{ngerman}±4270\textsuperscript{f}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
1050\selectlanguage{ngerman}±520\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
320\selectlanguage{ngerman}±95.0\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
65\selectlanguage{ngerman}±13\textsuperscript{g}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
500\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
2800\selectlanguage{ngerman}±1600\textsuperscript{f}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
12 400\selectlanguage{ngerman}±2790\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
60\selectlanguage{ngerman}±7.0\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
5000\selectlanguage{ngerman}±2640\textsuperscript{g}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
2500\selectlanguage{ngerman}±260\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
12 600\selectlanguage{ngerman}±11 000\textsuperscript{e}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
835\selectlanguage{ngerman}±480\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
25\selectlanguage{ngerman}±7.0\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
95\selectlanguage{ngerman}±20\textsuperscript{d}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
1000\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
7400\selectlanguage{ngerman}±3710\textsuperscript{e}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
11 200\selectlanguage{ngerman}±4000\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
40\selectlanguage{ngerman}±10\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
13 200\selectlanguage{ngerman}±2180\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
2300\selectlanguage{ngerman}±1130\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
17 100\selectlanguage{ngerman}±8440\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
440\selectlanguage{ngerman}±260\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
90\selectlanguage{ngerman}±40\textsuperscript{d}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Old leaves}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
Control\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
40\selectlanguage{ngerman}±20.0\textsuperscript{h}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
10 100\selectlanguage{ngerman}±4200\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
50\selectlanguage{ngerman}±20\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
10 500\selectlanguage{ngerman}±2770\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
4790\selectlanguage{ngerman}±1900\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
5490\selectlanguage{ngerman}±2740\textsuperscript{g}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
1640\selectlanguage{ngerman}±1270\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
15\selectlanguage{ngerman}±3.3\textsuperscript{h}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
200\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
1700\selectlanguage{ngerman}±1060\textsuperscript{g}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
8680\selectlanguage{ngerman}±1730\textsuperscript{e}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
34\selectlanguage{ngerman}±5.5\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
15 600\selectlanguage{ngerman}±12 200\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
3200\selectlanguage{ngerman}±2210\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
14 100\selectlanguage{ngerman}±12 600\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
1040\selectlanguage{ngerman}±740\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
55\selectlanguage{ngerman}±20\textsuperscript{g}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
500\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
9000\selectlanguage{ngerman}±5400\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
10 500\selectlanguage{ngerman}±2540\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
40\selectlanguage{ngerman}±8.0\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
12 200\selectlanguage{ngerman}±4720\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
2400\selectlanguage{ngerman}±620\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
21 800\selectlanguage{ngerman}±8850\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
430\selectlanguage{ngerman}±110\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
70\selectlanguage{ngerman}±27\textsuperscript{e}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
1000\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
10 700\selectlanguage{ngerman}±5370\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
8170\selectlanguage{ngerman}±1330e\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
45\selectlanguage{ngerman}±6.0\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
12 500\selectlanguage{ngerman}±3600\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
3090\selectlanguage{ngerman}±540\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
24 400\selectlanguage{ngerman}±10 700\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
630\selectlanguage{ngerman}±140\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
100\selectlanguage{ngerman}±60\textsuperscript{c}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textbf{Twigs}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
Control\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
8000\selectlanguage{ngerman}±220\textsuperscript{e}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
17 100\selectlanguage{ngerman}±1580\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
25\selectlanguage{ngerman}±3.60\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
8000\selectlanguage{ngerman}±220\textsuperscript{f}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
1460\selectlanguage{ngerman}±270\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
2900\selectlanguage{ngerman}±1300\textsuperscript{i}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
240\selectlanguage{ngerman}±49.0\textsuperscript{e}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
445\selectlanguage{ngerman}±200\textsuperscript{a}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
200\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
12 700\selectlanguage{ngerman}±4150\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
11 100\selectlanguage{ngerman}±2970\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
35\selectlanguage{ngerman}±5.50\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
7150\selectlanguage{ngerman}±3600\textsuperscript{f}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
2600\selectlanguage{ngerman}±1280\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
4300\selectlanguage{ngerman}±1220\textsuperscript{h}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
840\selectlanguage{ngerman}±603\textsuperscript{b}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
100\selectlanguage{ngerman}±34.0\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
130\selectlanguage{ngerman}±65.0\textsuperscript{c}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
500\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
8000\selectlanguage{ngerman}±520\textsuperscript{e}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
10 800\selectlanguage{ngerman}±3700\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
60\selectlanguage{ngerman}±8.02\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
7500\selectlanguage{ngerman}±4100\textsuperscript{f}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
2270\selectlanguage{ngerman}±430\textsuperscript{d}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
5000\selectlanguage{ngerman}±1150\textsuperscript{h}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
650\selectlanguage{ngerman}±235\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
60\selectlanguage{ngerman}±35\textsuperscript{g}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
1000\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
3100\selectlanguage{ngerman}±1700\textsuperscript{f}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
11 700\selectlanguage{ngerman}±3780\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
50\selectlanguage{ngerman}±10.4\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
8600\selectlanguage{ngerman}±3900\textsuperscript{e}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
2040\selectlanguage{ngerman}±790\textsuperscript{e}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
17 400\selectlanguage{ngerman}±6690\textsuperscript{c}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
1100\selectlanguage{ngerman}±840\textsuperscript{a}\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
\textless{}LOD\strut
\end{minipage} & \begin{minipage}[t]{0.07\columnwidth}\raggedright\strut
80\selectlanguage{ngerman}±33.0\textsuperscript{e}\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}\selectlanguage{ngerman}

\textless{}LOD = below Limit of Detection; LOD for Ni = 0.03 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1}, LOD for P = 6.00 \selectlanguage{greek}µ\selectlanguage{english}g g\textsuperscript{-1}

\textbf{Table 3.} pH, Diethylenetriaminepentaacetic acid (DTPA) and
Strontium nitrate (Sr(NO\textsubscript{3})\textsubscript{2}) extractable
concentrations in soils of different treatment levels of Mn of \emph{G.
bidwillii} at harvest and soil of \emph{G. acmenoides} . Values with
different small letters in the same column are significantly different
(\emph{p} \textless{}0.05).\selectlanguage{english}
\begin{longtable}[]{@{}llllllll@{}}
\toprule
Gossia bidwillii & \textbf{pH} & \selectlanguage{greek}\textbf{DTPA-extractable concentrations
(µg g\textsuperscript{-1})}\selectlanguage{english} & \selectlanguage{greek}\textbf{DTPA-extractable concentrations
(µg g\textsuperscript{-1})}\selectlanguage{english} & \selectlanguage{greek}\textbf{DTPA-extractable concentrations
(µg g\textsuperscript{-1})}\selectlanguage{english} &
\selectlanguage{greek}\textbf{Sr(NO\textsubscript{3})\textsubscript{2}-extractable
concentrations (µg g\textsuperscript{-1})}\selectlanguage{english} &
\selectlanguage{greek}\textbf{Sr(NO\textsubscript{3})\textsubscript{2}-extractable
concentrations (µg g\textsuperscript{-1})}\selectlanguage{english} &
\selectlanguage{greek}\textbf{Sr(NO\textsubscript{3})\textsubscript{2}-extractable
concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\tabularnewline
\midrule
\endhead
\selectlanguage{greek}\textbf{Treatment levels (µg g\textsuperscript{-1})}\selectlanguage{english} & & \textbf{Co} &
\textbf{Mn} & \textbf{Zn} & \textbf{Co} & \textbf{Mn} &
\textbf{Zn}\tabularnewline
Control & 5.95 & 1.60\selectlanguage{ngerman}±0.07\textsuperscript{c} & \textless{}LOD &
32.5\selectlanguage{ngerman}±4.24\textsuperscript{c} & 1.00\selectlanguage{ngerman}±0.01\textsuperscript{c} &
\textless{}LOD & 7.30\selectlanguage{ngerman}±1.24\textsuperscript{d}\tabularnewline
Mn 200 & 5.80 & 5.80\selectlanguage{ngerman}±0.30\textsuperscript{b} & \textless{}LOD &
44.5\selectlanguage{ngerman}±10.0\textsuperscript{c} & 1.20\selectlanguage{ngerman}±0.04\textsuperscript{c} & 6.52\selectlanguage{ngerman}±3.40
& 12.6\selectlanguage{ngerman}±6.70\textsuperscript{b}\tabularnewline
Mn 500 & 5.61 & 4.90\selectlanguage{ngerman}±0.75\textsuperscript{b} & 27.2\selectlanguage{ngerman}±6.03 &
60.0\selectlanguage{ngerman}±13.4\textsuperscript{b} & 3.45\selectlanguage{ngerman}±0.22\textsuperscript{b} & 10.1\selectlanguage{ngerman}±5.60
& 18.9\selectlanguage{ngerman}±3.40\textsuperscript{b}\tabularnewline
Mn 1000 & 5.27 & 12.7\selectlanguage{ngerman}±2.04\textsuperscript{a} & 37.30\selectlanguage{ngerman}±9.01 &
126\selectlanguage{ngerman}±56.07\textsuperscript{a} & 6.70\selectlanguage{ngerman}±1.90\textsuperscript{a} & 17.8\selectlanguage{ngerman}±8.01
& 30.3\selectlanguage{ngerman}±7.55\textsuperscript{a}\tabularnewline
Gossia acmenoides & \textbf{pH} & \selectlanguage{greek}\textbf{DTPA-extractable
concentrations (µg g\textsuperscript{-1})}\selectlanguage{english} & \selectlanguage{greek}\textbf{DTPA-extractable
concentrations (µg g\textsuperscript{-1})}\selectlanguage{english} & \selectlanguage{greek}\textbf{DTPA-extractable
concentrations (µg g\textsuperscript{-1})}\selectlanguage{english} &
\selectlanguage{greek}\textbf{Sr(NO\textsubscript{3})\textsubscript{2}-extractable
concentrations (µg g\textsuperscript{-1})}\selectlanguage{english} &
\selectlanguage{greek}\textbf{Sr(NO\textsubscript{3})\textsubscript{2}-extractable
concentrations (µg g\textsuperscript{-1})}\selectlanguage{english} &
\selectlanguage{greek}\textbf{Sr(NO\textsubscript{3})\textsubscript{2}-extractable
concentrations (µg g\textsuperscript{-1})}\selectlanguage{english}\tabularnewline
& & \textbf{Co} & \textbf{Mn} & \textbf{Zn} & \textbf{Co} & \textbf{Mn}
& \textbf{Zn}\tabularnewline
& 4.20 & 0.98 & 1580 & 12.2 & 0.19 & 1650 & 3.00\tabularnewline
\bottomrule
\end{longtable}

\textless{}LOD = below Limit of Detection; LOD for Mn = 0.07 \selectlanguage{greek}µ\selectlanguage{english}g
g\textsuperscript{-1}

\selectlanguage{english}
\FloatBarrier
\end{document}

