Accepted Manuscript

Lithium toxicity in plants: Reasons, mechanisms and remediation possibilities – A

review

Babar Shahzad, Mohsin Tanveer, Waseem Hassan, Adnan Noor Shah, Shakeel
Ahmad Anjum, Sardar Alam Cheema, Iftikhar Ali

PII: S0981-9428(16)30215-7
DOI: 10.1016/j.plaphy.2016.05.034
Reference: PLAPHY 4565

To appear in: Plant Physiology and Biochemistry

Received Date: 15 February 2016

Revised Date: 25 May 2016
Accepted Date: 25 May 2016

Please cite this article as: B. Shahzad, M. Tanveer, W. Hassan, A.N. Shah, S.A. Anjum, S.A. Cheema,
I. Ali, Lithium toxicity in plants: Reasons, mechanisms and remediation possibilities – A review, Plant
Physiology et Biochemistry (2016), doi: 10.1016/j.plaphy.2016.05.034.

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 ACCEPTED MANUSCRIPT

Lithium toxicity in plants: reasons, mechanisms and remediation possibilities -

A review

Babar Shahzad a, c, Mohsin Tanveerb, c, f*, Waseem Hassane, Adnan Noor Shahf, Shakeel
Ahmad Anjuma, Sardar Alam Cheemaa, Iftikhar Alia
a
Department of Agronomy, University of Agriculture Faisalabad, Pakistan

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b
School of Land and Food, University of Tasmania Hobart Australia
c
Equal contributed in manuscript as first author

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e
Department of Soil and Environmental Sciences, Muhammad Nawaz Shareef University
of Agriculture, Multan, Pakistan

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f
College of Plant Science and Technology, Huazhong Agricultural University Wuhan
China

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Corresponding author: Mohsin.tanveer@utas.edu.au
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Abstract
Lithium (Li) is a naturally occurring element; however, it is one of the non-essential
metals for life. Lithium is becoming a serious matter of discussion for the people who do
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research on trace metals and environmental toxicity in plants. Due to limited information
available regarding its mobility from soil to plants, the adverse effects of Li toxicity to
plants are still unclear. This article briefly discusses issues around Li, its role and its
essentiality in plants and research directions that may assist in inter-disciplinary studies to
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evaluate the importance of Li s toxicity. Further, potential remediation approaches will

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also be highlighted in this review. Briefly, Li influenced the growth of plants in both
stimulation and reduction ways, depending on the concentration of Li in growth medium.
On the negative side, Li reduces the plant growth by interrupting numerous physiological
processes and altering metabolism in plant. The contamination of soil by Li is becoming
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a serious problem, which might be a threat for crop production in the near future.
Additionally, lack of considerable information about the tolerance mechanisms of plants
further intensifies the situation. Therefore, future research should emphasize in finding
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prominent and approachable solutions to minimize the entry of Li from its sources
(especially from Li batteries) into the soil and food chain.
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Keywords: Lithium, Effects and mobility, Plant growth, Physiology, Research

perspectives

Capsule abstract:
Li not only reduces but also stimulates plant growth, depending on its concentration in
growth medium.
Li interferes with numerous physiological and metabolic processes to influence plant
growth.
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1. Introduction

1.1. Sources of Li in environment

Lithium (Li) exists in its elemental form and is highly reactive as a pure element. Due to
its highly reactive behavior, Li reacts with other elements and exists as salts or minerals
(Bleiwas and Coffman, 1986). Lithium minerals are mined around the world, e.g., in

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Zaire, Zimbabwe, Western Australia, Canada, Russia and China to make various lithium
mineral products (Moore, 2007). Lithium added into the ecosystem by various sources
including industrial waste water and nearby water resources near to Li-related industry

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(Hull et al., 2014), disposal of Li batteries (Al-Thyabat et al., 2013) and Li brines
(Shacklette and Boerngen, 1984; Hem, 1992). Among these, Li brines are a major source

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of Li introduction (Table 1). Different other studies on other hand, indicated that Li is
present in higher concentrations than typically encountered in natural surface waters (e.g.
Hem, 1985) and was at concentrations similar to those documented due to mineralization

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or mining related enrichment (Kszos and Stewart, 2003; Jones et al., 2013). Lithium (Li)
is an alkali metal and is widely distributed in the environment. Furthermore, Li is the 27th
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most abundant element and covers about 0.006% (wt) of the earth crust (Habashi, 1997).
In water environments, Li compounds are highly soluble and relatively inert in dissolved
form, have been used as conservative traces in different hydrological studies (Tate et al.,
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1995; Stewart and Kszos, 1996; Fouillac and Michard, 1981), while a collaborative study
of Department of Energy, US and Jeppson (1978) reported that Li reacts readily with
water to form hydrogen gas (a hazard during some accidents).
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1.2. Essential nature of Li

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It is unclear whether Li is essential for plant growth while some studies reported
different essential characters of Li in different living organisms. Different plant species
can absorb considerable concentration of Li, though it seems not to be essential for the
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growth and development (Jiang et al., 2014). Contrasting evidence reveals that Li can
have some important functions in the metabolism in halophytes. However, Li is toxic to
plants when present in high concentration in soils (Kabata-Pendias and Mukherjee,
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2007). Mulkey (2005) found that Li alters maize root growth and gravitropic response
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under higher Li concentration. Some studies reported that Li inhibits the inositol cycle
and calcium signaling in plants (Berridge, 1993; Naranjo et al., 2003). Further
considerable modifications were observed in the metablomes and transcriptomes of
Brassica carinata seedlings under Li stress, which suggested that Li plays an essential
role in the biochemistry of plants (Li et al., 2009). Low levels of Li conversely have also
improved plant productivity in some instances, increasing yield, hastening maturation,
and increasing resistance to disease (Bradford, 1966). For instance, Li at low
concentration (5 mg dm−3) effectively increased maize biomass production by 15%
(Hawrylak-Nowak et al., 2012). Hence, yet there is no clear evidence that can prove that
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Li is an essential or beneficial element for plants. Therefore, the essentiality of Li for

plant is still disputable.

The essential nature of Li in animals, including humans is also questionable for

researchers however; Li deficiency caused anomalies such as reduced lactation and
reproduction in goats (Mikhaleva et al., 2001) and rats (Pickett, 1983; Schrauzer, 2002).

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Low Li intakes result in an altered negative behavior and aggressiveness in humans
(Dawson et al., 1970, 1972) and in rats (Klemfuss and Schrauzer, 1995). In another study,
it was observed that when rats were fed a Li-deficient diet (0.0066 g Li g-1 diet) showed

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suppressed level-press avoidance behavior as compared to animals provided a Li-fortified
diet (0.11 Li g/g diet) (Ono and Wada, 1991). Moreover, Klemfuss and Schrauzer (1995)
and Schrauzer (2002) documented a role of Li in some biological processes such as

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enzymatic activity, transporters and channels activity, energy production and/or circadian
movement in living organism, which further emphasizes the importance to study Li in
living organisms. In animals, Li is non-bio accumulative; however their compounds have

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toxic effects and those effects are Li dose-dependent and animal species dependent.
Kszos et al. (2003) found that Li even at low concentration (0.5 mg L-1) reduced
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reproduction of fathead minnow growth and Ceriodaphnia. Aral and Vecchrio-Sadus
(2008) reported that Li concentration at 526-840 mg kg-1 are lethal to rats, while at
concentration of more than 20 mg L-1 Li or higher there is a risk of death due to Li
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toxicity.

Many biological functions of Li are probably due to its small ionic radius. This
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induces powerful polarizing effects and may displace K+, Mg2+ and Ca2+ at various
biological membranes and enzyme binding sites (Schrauzer, 2002). Duff et al. (2014)
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reported that Li+ might play a larger role in the inhibition of Mg2+-containing proteins in
plant as well as mammalian systems. The essential nature of Li is not fully explored.
Nonetheless, it can be suggested that Li can have similar functions with Mg or Ca in
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plants or may replace them during normal metabolic processes in plants.

1.3. Importance of Li in plants

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Compounds of Li are widely distributed in nature, but usually occur only in

minute concentrations. It is reported that average values for Li compounds are 30 mg kg-1
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in the earth's crust, 25 mg kg-1 in soil, 2 mg kg-1 in drinking water, 170-190 µg L-1 in
seawater and 2 ng m-3 in the atmosphere (Birch, 1988; Ribas, 1991). Lithium
concentration varies across different water bodies (e.g. fresh water, lakes, sea water,
ground water), with indicative values in Table 2, is further highlighting the importance to
study Li in plant. In foodstuffs, Li concentration varied widely ranged from 0.01 ppm in
bananas to 55 ppm in oats (Shacklette et al., 1978). Industrial use of Li includes
industries producing colorings, batteries and metal alloys manufacturing industries, and
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Li based pool-cleaning chemicals, which are major sources of Li contamination for water
bodies and soil, thus polluting environments and providing a direct source of Li toxicity
for plants (Schrauzer, 2002). In the wetlands of Nevada, U.S.A., Li at concentrations
greater than 1 mg L-1 have been noted (Hallock and Hallock, 1993). Bradford (1963)
found reduced and stunted growth of citrus in southern California, U.S.A., with the use of
highly Li-contaminated water for irrigation. He also recorded that the Li concentration in

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100 out of 400 water samples was more than 0.05 mg L-1, which may cause toxic and
undesirable effects in citrus plants. Lithium concentrations in different soil series has
been documented that ultimately reduced food quality (Table 3). Threshold

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concentrations of Li in plants are highly variable, and moderate to severe toxic effects at
4 to 40 mg Li kg-1 was observed in citrus leaves (Kabata-Pendias and Pendias, 1992).
Borovik-Romanova (1965) analyzed 138 plants from eight soil types and found that

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plants generally have an average content of 0.15 to 0.3 mg Li kg-1, but plants rooted in
saline soils may contain two to ten times more contents than that level. Robinson et al.
(1971) found an average of 0.3 to 0.42 mg Li kg-1 (dry matter) in various classes of

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vegetation. Water-soluble forms of Li in the soil profile reached about 5% of the total soil
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contents and expected to increase in areas due to more Li contents in rocks and soils
(Kabata-Pendias and Pendias, 1992). In addition, the abundance of Li in surface soils
seemed to have a correlation with soil units (Kabata-Pendias and Pendias, 1992). Grand
means for Li varied from 1.3 mg kg-1 in light organic soils to 56 mg kg-1 in calcareous
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soils. Daily intake of Li from different foodstuffs (Table 4) provoked the study of Li toxic
effects in food, which subsequently affects different metabolic processes in plants and
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other living organisms. Therefore, the current review compiled previous information
regarding Li toxicity in plants to underpin the mechanisms behind Li toxicity.
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2. Li uptake, translocation and accumulation in plants

Weathering process involves the oxidation and results in the release of Li from parental
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material to clay soil where Li is fixed by organic matter or by some other mineral oxides
and ultimately accumulates in phosphate rocks. On the other hand, Li accumulation in
soil could also be influenced by soil conditions, clay contents or initial contents of Li in
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parental material (Kabata-Pendias and Pendias 1992). Anderson et al. (1988) found a
positive correlation between soil clay contents and total and exchangeable Li contents in
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soil but observed an inverse relation between clay contents and water-soluble Li. They
also found a weak, but significant correlation (r=0.623) between clay and exchangeable
Li in Piedmont soils. Li is available in trace amount in soil and Li contents ranged from
0.002 to 63 ppm in various soils (Vergara-Edwards et al., 1986), while Schruazer (2002)
observed Li contents ranging from 7 to 200 µg g-1 in clay fractions. Siroonian et al.
(1959) considered five possible means by which minor elements may associate with
minerals; in mechanically admixed impurities, in inclusions and alteration products, in
structural voids or cavities, in an adsorbed phase, and in solid solutions.
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Lithium is widely distributed so one could expect that plants can easily uptake Li.
Furthermore, though Li is considered as a non-essential and non-physiological element.
However, almost all plants can absorb Li from their immediate environment.
Nonetheless, variation exists among plant species in Li uptake and accumulation. Some
plants are hyper or bio-accumulators of Li (Li et al., 2009), while others showed Li
uptake below a threshold level and further increase in Li resulted in detrimental growth

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reduction (Hawrylak-Nowak et al., 2012). The exact mechanism of Li uptake and
translocation within plant is yet unclear, although some studies suggested that Li has
physiochemical properties similar to other alkali metals such as Na or K and thus can be

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absorbed by the same mechanism used for these elements (Shkolnik, 1984).

Lithium may share the same pathways and the same transport carrier across cell

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membranes as K+ does and may therefore, easily be taken up and transported in leaves
(Kabata-Pendias and Mukherjee, 2007). Some plants showed a competitive interaction
between Ca and Li in absorption by plant roots. Thus, significant levels of Li in soils,

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then, could potentially influence Ca uptake and cycling (Epstein, 1960). Further, different
plant species also showed different absorbing and tolerance capabilities under Li stress.
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Hawrylak-Nowak et al. (2012) observed that increases in Li concentration in growth
medium caused a dose-dependent increase in Li uptake in maize and sunflower shoot, but
sunflower (3292 kg−1 dry weight) exhibited higher accumulation of Li as compared to
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maize (695 mg Li kg−1 DW) when supplemented by Li at the concentration of 50 mg Li

dm−3 in the nutrient solution. Further, Arabidopsis plants when exposed to LiCl in
hydroponic culture showed curling of leaves, and chlorosis mainly in older leaves (Duff
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et al., 2014). Similarly, in tobacco leaves, necrotic lesions due to Li were more obvious in
older leaves (Naranjo et al., 2003).
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The uptake, translocation and accumulation of Li by different plant species could

also be partially explained by different experimental conditions, plant ages, sampling date
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and exposure. Therefore, exact information regarding Li uptake and translocation in

plants could not be interfered. A significant increase of Li uptake and translocation in the
above ground parts of agricultural crops were observed under soil application of Li
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(Jurkowska et al., 1991 and 1995), while Kalinowska et al. (2013) observed less Li
accumulation in shoot than root and suggested that this difference might be due to the Li
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distribution pattern between below-ground and above-ground parts of plants when grown
in nutrient solution and can be different from those of plants grown in the soil. In another
study, Magalhães et al. (1990) documented that there was significant difference in the
accumulation of Li among different hydroponically grown dicotyledonous plants and
found that radish and watercress accumulated more Li in tissues as compared to lettuce.
Li et al. (2009) used inductively coupled plasma mass spectrometry to examine the
pattern and behavior of different Brassica carinata seedlings types for the accumulation
of Li. They also observed significantly dose-dependent difference among these seedling
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types. They noted that initially yellow and brown-seeded lines avoided Li uptake and
then showed accumulation of equal amount of Li under low concentration, while under
high concentration (≤ 60 mM), Li accumulation was compromised in yellow seeded lines
only. Brown seeded lines accumulated more than 80,000 ppm (dry weight) Li at the same
high concentration. Moreover, a number of factors relating to soil such as pH or moisture
contents, could also influence the availability, uptake and accumulation of Li in plants.

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Some plants of the Solanaceae family, when grown in an acidic climatic zone accumulate
more than 1,000 mg kg-1 Li. The highest uptake of Li was noted in those plant species
growing on solonetz and solonchak soils or other soils having increased contents of alkali

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metals (Kabata-Pendias and Pendias, 1992). Aral and Vecchio-Sadus (2008) documented
that generally Li in higher concentrations is toxic to all plants and more Li is taken up
from acidic soil as compared to alkaline soils. This is probably because soil acidity

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corresponds to an increase in the solubility of Li and plant Li levels are correlated with
the concentration of these elements in soil (Schrauzer, 2002; Aral and Vecchio-Sadus,
2008). At lower soil pH, there is a reduced negative surface charge, resulting in lower

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binding sites available for cations. The higher solubility of the elements will result in
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greater plant uptake of these elements, which will subsequently result in greater plant
uptake of Li

Li does not translocate to younger or other plant organs as higher concentration of

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Li in older leaves and immobile movement of Li in phloem tissue of wheat was noted
(Zeller and Fuller, 2000). While some studies indicated that Li is an immobile element
and its toxic symptoms appear in older leaves but studies also corroborated that Li shows
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movements similar to K or Ca. Mason and Maskell (1931) observed that K was not
accumulated in cotton leaves but was re-exported from leaves to roots where it might
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retard the rate of salt absorption. However at the same time, there was no backward
movement of Ca and was unable to move within phloem. It can be suggested from this
study that K can be cut off from plant without adverse effects. New leaves can get K from
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phloem transport or by re-export from older leaves, while this mechanism does not work
for Ca; therefore, the need for Ca remained throughout the development of plant and can
only be acquired directly from roots rather than through re-export. Therefore, Li
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resembles Ca in not being re-exported from the leaf. This behavior may be due to failure
of mobilization in the leaf or due to the incapacity of Li to move in the phloem. Birch-
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Hirschfeld (1920) found that the rate of movement of Li through the phloem was small. A
similar mechanism was also observed in wheat when exposed to different Li treatments
with different sampling times (Kent, 1941). Kent concluded that Li bears more
resemblance to Ca than to the other alkali metals as he observed that after another few
days of Li treatment, young wheat seedlings showed Li accumulation in roots and then
after another few days, a high amount of Li was found in older leaves as compared to
roots, suggesting that Li is an immobile element. On the other hand, he also observed loss
of Li from the roots to the soil after transplanting from one pot to another pot, suggesting
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that the movement of Li in and out of the plant is governed principally by the direction of
the Li gradient, this redistribution is hindered by the immobility of Li in the first wheat
leaf. In a hydroponic experiment, Li concentration in nutrient solution to 12 ppm,
increased cucumber fruit yield, fruit sugar, and ascorbic acid levels, but Li did not
accumulate in the fruit (Rusin, 1979). In a field experiment in soil ranging from 1.4 to 2.4
ppm Li, addition of 1 kg Li ha-1 increased potato tuber citric and malic acids (Vlasyuk, et

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al., 1973).

3. Effect of Li toxicity in plants

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3.1. Effects on growth and development

Plant growth is highly governed by a number of factors including weather

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conditions, genetic factors, topography, and soil fertility. A number of metals are
important for the growth of plants. However their essentiality depends on their
concentrations in plant and environment (Anjum et al., 2015a). Furthermore, some metals

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are toxic to plants and an increase in their concentration from optimum level, however,
retards plant growth and yield (Anjum et al., 2015b; 2016a; 2016b). Although, the
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essential nature and toxic effects of Li on higher plants are not yet clear, previous
evidences indicates that Li induces considerable reduction in plant growth as the Li-salts
are highly toxic and trigger formation of necrotic regions (Naranjo et al., 2003).
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Nonetheless, different plant species showed plastic behavior in sensitivity and tolerance
to Li toxicity. Schrauzer (2002) and Kabata-Pendias and Mukherjee (2007) noted that
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plants of Asteraceae and Solanaceae families showed tolerance against Li toxicity and
exhibited normal plant growth while Aral and Vecchio-Sadus (2008) found that citrus
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plants are sensitive to Li stress. Other Li toxic effects included altered rhythmic
movement of petals (Birch, 1991) and disrupted pollen development (Zonia and Tupy,
1995).
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Plant roots are the first organ that comes in contact with Li in soil and Li in excess
alters root gravitropic growth of maize roots (Mulkey, 2005). Furthermore, in spinach, Li
toxicity affects cold-induced de-phosphorylation of microtubules in mesophyll cells of
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spinach (Bartolo and Carter, 1992). In the Li-rich soils, damage of root tips and chlorotic
and necrotic spots on leaves have been observed in corn (Kabata-Pendias and Mukherjee,
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2007). Necrotic spots are typical for incompatible (avirulent) plant–pathogen interactions,
and the induction of necrotic spots in Li-treated plants seems to be mediated by ethylene
(Naranjo et al., 2003). Jurkowska et al. (1998) noted considerable decrease in the yield of
oat exposed to 25 mg Li kg−1, and maize and spinach, when exposed to 40 mg Li kg−1
cultivated in the soil. In lily pollen, 50mM Li can inhibit enzyme activities by 50%
(Gumber et al., 1984).
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The reduction of plant growth depends on Li concentration and plant species

(Table 5). Lithium at low concentration stimulates plant growth, while at higher
concentrations Li can completely inhibit plant growth. Moreover, Li toxicity-induced
symptoms are not distinct and are difficult to recognize; however, Li-injured broad-leaf
plants generally exhibit necrosis along the leaf margins, subsequent interveinal chlorosis
and leaf abscission. Kalinowska et al. (2013) found that addition of Li at low rates up to

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20 mg Li dm−3 had a positive and stimulatory effect on the growth and elongation of the
root and shoot of lettuce, while at 50 mg Li dm−3, Li significantly reduced root and shoot
biomass production, and a further increase in Li concentration to 100 mg Li dm−3 induced

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complete growth inhibition. Apparent symptoms of Li toxicity such as necrotic spots on
older leaves of lettuce clearly appeared upon exposure to 100 mg Li dm−3 (Fig. 1a)
(Kalinowska et al., 2013). Hawrylak-Nowak et al. (2012) studied the toxic effects of Li

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on the growth of maize and sunflower and observed that at low concentration of Li
growth of both plants was stimulated; however, at higher supplies the assimilating organ
area of sunflower and maize plants decreased by 27% and 31% respectively with

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increasing concentrations of Li. Necrotic spots on sunflower leaves were also observed at
a supply of 50 mg Li dm−3 (Fig. 1b). Furthermore, they also noted that exposure of maize
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plants to 25 mg Li dm−3 resulted in the reduction of leaf area by 29% as compared to a
control. Bingham et al. (1964) examined Li toxicity in a number of plant species and
divided those species on the basis of Li concentration in tissues at 25% growth
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depression, and /or full growth inhibition. He also indicated the range of Li
concentrations that induced no toxic, mildly toxic and highly toxic effects (Table 6). In
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conclusion, Li toxicity varies among plant species. In Phaseolus and Helianthus,

circumnutation, but not growth, was affected by persistent Li application (Zachariassen
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and Johnsson, 1988; Millet and Badot, 1996). Lithium also affected movement of Cassia
fasciculata leaves (Gaillochet, 1981) and Desmodium motorium lateral leaflets (Weber et
al., 1992) and caused changes to the gravi-responsiveness of roots in Pisum sativum
(Belyavskaya, 2001). Stevenson et al. (2000) suggested that Li+ acts via the inositol
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signaling pathway and is also involved in glutamate-receptor-, and Ca2+- dependent plant
responses and these processes together with H+-ATPase are involved in growth,
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circumnutation and excitability. Furthermore, it has also been demonstrated that Li

affects plant metabolism at a membrane level; however, its role seems to be non-specific
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as it substitutes for other monovalent cations in plant cells (Kabata-Pendias and

Mukherjee, 2007). Allender et al. (1997) suggested that adverse effects of Li on
hydroponically grown cotton might be because the Li+ ion readily crosses cell
membranes and interferes with calcium metabolism.

Reduction in water content is another toxic effect of Li. It reduces tissue turgidity
and growth. However, contradictions are also documented in this regard, e.g., lettuce
shoot and roots exhibited higher water content when exposed to low concentration of
LiCl or LiOH as compared with higher concentrations of Li (Kalinowska et al., 2013).
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Magalhães et al. (1990) concluded that reduced dry weight for roots and leaves of radish
and leaves of watercress under Li stress was due to reduced water retention in tissues.
Duff et al. (2014) substantiated that finding when plants of Arabidopsis were exposed to
Li+ for a period of 3 weeks in that low values of relative water content and structural
independent pigment index were observed. In wheat, Li reduced net water content and
dry weight; however, it was not clear whether tissue desiccation under Li stress was due

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to reduced water uptake or elevated transpiration rate (Kent, 1941). Contrarily, Vlasyuk
et al. (1979) suggested the mechanism behind increased water absorbance under low Li
concentration. They proposed that low Li concentrations increased the amount of water

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adsorbed to macromolecules, but at high concentration, Li competed for water and acted
as a dehydrating agent. Evidence supporting this hypothesis may come from the work of
Hassan (1954), who stated that the large degree of hydration of Li+ ions is related to

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changes in cytoplasmic structure associated with increases in the size of radish root
cortical cells after treatment with Li. Lithium was also found to specifically perturb the
stacking equilibria in dinucleotide phosphates and polynucleotides. Based on this

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evidence, it was concluded that these effects resulted from a change in the hydration shell
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of the nucleic acids in the presence of Li (Powell and Richards, 1972).

In Brassica carrinata, Li et al. (2009) observed no significant toxic effect of Li up

to the 200 mg Li dm−3 on seed germination and growth related traits. However, above
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this concentration Li caused an abrupt decline in growth and ultimately complete growth
inhibition. They also noted that this reduction was associated with decreased chlorophyll
content and phenolic compound production under Li stress. In spinach and mustard
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plants, Li addition to the soil also increased plant biomasses, but only under conditions of
reduced light levels (Makus et al., 2006). Possibly, Li can act as a cofactor for the
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activation of some enzymes or may appear as a replacement factor to continue

metabolism due to its affinity to enzymes activated by Ca or Mg. Therefore, the
beneficial and toxic effects of Li still need further quantification to better resolve the
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dose-dependent relationships.

3.2. Effects on photosynthetic pigments and carbon assimilation

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Lithium is a non-physiological element that can induce toxic effects on

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photosynthetic pigments and carbon assimilation. Hawrylak-Nowak et al. (2012)

observed that supplementation of Li at a concentration of 5 to 50 mg dm−3 in a nutrient
solution did not influence chlorophyll a and b contents in sunflower, while carotenoid
contents were considerably decreased by 16% at a concentration of 25 mg Li dm−3.
Contrary to this result, maize showed a reduction in chlorophyll a and b contents by about
45% and carotenoid contents by 67%, with the appearance of necrotic spots on leaves,
under a high Li concentration of 50 mg dm−3 as compared with a control (Fig. 1c)
(Hawrylak-Nowak et al., 2012). Li et al. (2009) noted that Li salt-stress depressed
chlorophyll content in different B. carinata seedlings. This depression could be related to
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changes observed in several other plant species under different forms of stress, e.g., the
disintegration of chloroplast fine structure, instability of pigment protein complexes, or
changes in the quantity and composition of metabolites (Dubey, 2005).

Lithium causes development of necrotic regions, as reported for leaves of lettuce

(Kalinowska et al., 2013), and red kidney bean, sour orange and tomato (Bingham et al.

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1964). The development of necrotic regions could be related to the breakdown of
chlorophyll contents in leaves. Another possible reason for the loss of chlorophyll from
necrotic regions might the increased catalytic action of chlorophyllase, pheophorbide

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oxygenase, red chlorophyll catabolite reductase, and Mg-dechelatase, which are
responsible for chlorophyll bleaching (Harpaz-Saad et al., 2007). It has been reported that
Li is a non-essential element to plants, although there is some evidence of its effects on

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photosynthesis, synthesis and translocation of sugars, a great number of enzymatic
processes and nitrogen metabolism (Shkolnik, 1984). Little information is available
regarding toxic effects of Li on photosynthesis and photosynthetic pigments. We

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speculate that some reasons for reduced photosynthetic activity in the presence of Li, may
be that Li induces damage to chloroplastic ultrastructure, inhibits electron chain
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reactions, substitutes for Mg in chlorophyll, reduces stomatal conductance, inhibits
Calvin cycle or causes the breakdown of plastoquinone and carotenoid. Future research
should focus on examining the effects of Li on these aspects to determine the mechanism
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behind Li-induced reduction of photosynthesis.

3.3. Effects on the uptake and accumulation of other mineral elements

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The essential nature of Li and its functions are not fully clear. Lithium belongs to
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group IA of the periodic table. It is an alkali metal with properties similar to other
members of group IA and some members of group IIA. Li has high ability to replace Na+
and K+ as well as Mg2+ and Ca2+ due to its smaller radius and high polarizing strength. Its
complexion properties are stronger than those of Na+ and K+, but weaker than those of
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Mg2+. Li can act as a substitute for K+ or Mg2+, and can bind to sites not occupied by K+
or Mg2+. Lithium can substitute for 50% of the K in plants (Codina et al., 1983). Birch
(1988) suggested that all alkali metal ions are exchanged more than 1000 times more
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rapidly than Mg2+; this may explain why Li preferably affects the activity of Mg2+
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containing enzymes. Magalhães et al. (1990) observed that Li increased iron uptake only
in lettuce, but did not increase it in other plants. Similarly, Li influenced K uptake by the
roots of radish while, in watercress, Li application only influenced N+ and Ca2+ uptake.

Kabata-Pendias and Mukherjee (2007) documented that Li interacts with Na+ and K+ and
with those enzymes that require Mg2+. Hawrylak-Nowak et al. (2012) observed no
substantial difference in the K+ uptake between control and Li-stressed maize plants,
while sunflower plants showed increased K+ uptake when exposed to a Li concentration
of 50 mg Li dm−3. Naranjo et al. (2003) conversely showed an abrupt depletion of K+
 ACCEPTED MANUSCRIPT

from tobacco leaves due to Li. However, this decrease in K+ contents did not correlate
with the intensity of visual Li-induced toxicity symptoms. The interaction of Li with K
and Ca is unclear; however, Jacobson et al. (1960) reported that Ca inhibits the
absorption of Li and normalizes K uptake under Li stress, suggesting that Li and Ca may
share the same absorption site in roots, while Li interferes with K uptake. Epstein (1960)
concluded that Li and Ca shared binding sites after he showed that the uptake of these

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two ions was competitive, as the sites transporting Li are distinct from those sites that
transport K. Under field conditions, Ca application was partially effective in ameliorating
Li toxicity in bean (Phaseolus vulagris) (McStay, 1980). In conclusion, Li showed some

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potency in influencing plant mineral nutrition, but very little and only older information
is available. Therefore, research is required to quantify the dose dependent relationship of
Li on the dynamics and mobilization of other nutrients in soil and plant tissues.

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3.4. Li induced oxidative damage and antioxidant system

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Even though studies reported that Li can stimulate growth at low concentration in
some plant species, Li in high concentration also reduces or in severe condition inhibits
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growth. The exact mechanism behind this is still unclear; however, one could suggest that
some harmful effects of Li could be related to oxidative stress (Kielczykowska et al.,
2004; Efrati et al., 2005; Oktem et al., 2005). Some studies indicated that the oxidative
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nature of Li is mediated by enhanced levels of lipid peroxidation (Tandon et al., 1998).

The enhanced oxidation in the presence of Li might be that it can inhibit the activation of
antioxidant enzymes or the transcription of genes responsible for antioxidant production.
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Furthermore, Li toxicity can also lead to higher intracellular levels of O2- or OH- radical
through a Fenton-type reaction known to initiate lipid peroxidation. Reactive oxygen
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species (ROS) react with numerous biological membranes, cellular organelles, and
biomolecules like proteins, lipids, photosynthetic pigments, nucleic acids (DNA or
RNA). The ultimate effect would be disruption of normal cellular functions and cell
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metabolism (Clemens, 2006; Singh et al., 2010). Unsaturated fatty acids are major
components of cell membranes. ROS production under Li toxicity can react with
aldehydes and replace hydrogen from unsaturated fatty acids and thus imparts cell
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membranes by distorting the lipid bilayer; however, the exact mechanism is still unclear.
Bhattacharjee (2005) documented that ROS reacts with bisallylic hydrogens present in
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polyunsaturated fatty acids by (I) initiating lipid radical formation, (II) progressing lipid
peroxyl radical formation as a product of lipid radical + oxygen reaction, and (III)
terminating production of non-radical products after molecular reactions with lipid
peroxyl radicals. As a result, ROS further disrupts lipid membranes and alters the
structures of various cell organelles including mitochondria, peroxisomes, and
chloroplasts (Małecka et al., 2008).

Antioxidant production is an adaptive mechanism in plants to scavenge ROS

produced under oxidative stress. In severe conditions ROS can inhibit the production of
 ACCEPTED MANUSCRIPT

enzymes such as superoxide dismutase (SOD), glutathione peroxidase (GPX) or catalase

(CAT) (Richard et al., 1997; Erakovic et al., 2000). Though little information is available
about oxidative damage due to Li toxicity, Nciri et al. (2012) observed in the kidney and
liver of rat that GPX activity was inhibited at 1mM Li, but at the same time SOD activity
was increased affirming that Li caused the production of superoxide ions (Nciri et al.,
2012). Hawrylak-Nowak et al. (2012) noted increased levels of lipid peroxidation as

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determined by the production of melanoaldehyde (MDA) in the leaves and roots of
sunflower and maize plants grown in the presence of 50 mg Li dm−3. This result suggests
the disturbance of membrane integrity while there was a non-significant effect of Li on

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MDA production in the roots of sunflower. Elevated MDA is generally positively
correlated with a decline in plant biomass. This could be because of oxidative stress,
resulting from an excessive formation of ROS, and is one of the important mechanisms of

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the toxic effects of Li ions; however, changes in the MDA content seem to be fully
ambiguous and are organ-dependent and species-specific. L-Ascorbic acid is another
important anti-oxidant compound. It is also known as vitamin C and is involved in the

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biosynthesis of neurotransmitters and collagen (Iqbal et al., 2004). L-Ascorbic acid along
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with glutathione make a non-enzymatic anti-oxidant defense system (Eraslan et al.,
2007). This compound is a scavenger of ROS created by Li stress. However, Kalinowska
et al. (2013) measured L-ascorbic acid accumulation in the leaves of lettuce plants and
found that its accumulation remained unaffected by different Li stress levels; however,
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there was a tendency for a decrease by 10–13% in L-ascorbic acid contents under the
highest Li concentration of 100 mg Li dm−3, but this reduction was not significant. Thus,
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they concluded that Li toxicity in plants is not related to the disturbances in the
accumulation of L-ascorbic acid in the leaf tissues.
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3.5. Effects on nucleic acids, enzymes and other processes

Nucleic acid metabolism is one of most important physiological processes in

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plants. It involves biosynthesis of nucleic acid, which contains phosphodiester backbone.

These phosphate groups possess the strong ability to bind with different cations. Li in
high concentration may impose detrimental effects on nucleic acid metabolism such as
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inhibiting DNA condensation, biosynthesis of protein, amides, or induce conformational

alteration in DNA; however, these effects still need to be studied further. Preliminary
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studies reported some toxic effects of Li on DNA, RNA and protein biosynthesis
pathways. Weeks (1956) reported that DNA condensation is partially governed by Na/K
ratio; however, Li competes with K and thus binds more readily and causes alteration in
the conformation of DNA. Vlasyuk et al. (1979) reviewed some studies and suggested
that Li in high concentration interrupts protein and nucleic acid metabolism. Evidence
from several studies showed direct toxic effects of Li on expression of some genes by
influencing transcription and translation (Vlasyuk and Kuz'menko, 1975; Vlasyuk et al.
1975a; Vlasyuk et al. 1975b). They also found that Li has a primary effect on the stability
 ACCEPTED MANUSCRIPT

and conformational state of DNA, and this in turn affects the metabolism of RNA and
proteins. Nonetheless, we have seen that Li may have both selective and non-selective
effects. The selective effects might be because of its physio-chemical properties, while
the non-selective effects might be due to its monovalent nature. Furthermore, it is evident
that the inhibitory and stimulatory effects of Li on plant growth depend on Li
concentration. Therefore, results and conclusions about the toxic effects and behavior of

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Li on nucleic acid and other genetic biomolecules cannot be interpreted or suggested.
Other effects of Li toxicity include abnormal cell division in the root apical meristem of
maize (Edwards, 1941). He also noted that Li at high concentration distorts metaphase

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chromosome arrangement, prevents cell division, and causes clumping of chromatin.
However, these results are apparently opposing the views about the effects of Li on DNA
and cell replication. On the other hand, the adverse responses may be secondary

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manifestations of some primary effects elsewhere in the system.

Li toxicity also influences normal functioning of numerous enzymes by binding to

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active sites. Results of Stracher (1960) showed that Li induces loosening of enzyme
conformation and results in exposure of reactive groups. Another hypothesis proposed
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that the mechanism of action of monovalent cations (e.g. Li+) on enzymes is that the
conformation of a monovalent cation-activated enzyme is flexible and depends upon the
cationic environment. The majority of monovalent cation-activated enzymes function in
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environments containing K. When these enzymes are in an environment where the

relatively large cation Li+ predominates, conformations may be assumed that are not
appropriate for catalysis. It was also suggested that the conformation of certain enzymes
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might be less specific, and less critical, and in these cases activation by Li would be
possible.
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It is apparent that, although hydrated ionic radii seem to contribute most greatly to
enzyme specificity, there is a need to consider the chemical properties of the ions
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involved. Some studies suggested a model to explain the interference of Li with K and its
concomitant effects (Thellier and Desbiez, 1977). They also reported that K is required
for the biosynthesis of a few K-ion specific proteins, but Li moved through an efficient
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pump with little Li leakage and concomitantly reduced the K content in cells. This could
be the way that Li could limit the biosynthesis of some K-ion specific proteins or also by
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affecting the protein biosynthetic apparatus (Carlier and Thellier, 1979, Kergosien et al.,
1979). With sycamore (Acer pseudoplatanus), Thellier (1980) observed that Li did not
cause a direct deactivation of the solute uptake machinery during transport. Instead, Li
prevented the establishment of the solute uptake mechanism by somehow inhibiting
transcription and translation. Thus, the effects of Li were thought to be primarily at the
level of protein biosynthesis. Lithium toxicity induced numerous disruptions in
morphogenetic traits of plants by interfering with the cell replication process or cell
polarity. Johnson (1975) found that Li does not bind strongly with protein. Due to rapid
 ACCEPTED MANUSCRIPT

Li exchange, protein structure would not be seriously altered even when bound at high
concentrations. However, there are reports showing the specific bonding ability of Li+
ions to amides (Bello et al., 1966), peptides (Armbruster and Pullman, 1974), and lipids
(Williams, 1973). The fact that Li forms these complexes may influence the ability of
proteins to react.

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Other detrimental effects of Li have also been reported such as increased
respiration process at wounded sites of Cichorium intybus (Laties, 1959); reduction in
floral induction and development due to toxic effects on phytochrome activity in

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duckweed (Kandeler, 1970); disruption of the circadian movement of petals in
Kalanachoe blossfeldiana (Englemann, 1972, 1973); modification in thigmo-
morphogenetic characteristics of Bidens pilosur and Cichorium intybus (Laties, 1959,

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Desbiez and Thellier, 1977, 1978); limitation of stomatal conductance and aging process
in geranium (Wortley, 1936, Carlier and Thellier, 1979); and elevation of stomatal
diffusion resistance in Phaseolus vulgaris plus inhibition of the seismonastic closure of

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leaflets of Mimosa pudica (McStay, 1980).
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Lithium in high concentration induced a number of alterations and inevitable
modification in whole plant biological processes; however, it is not wise and even
possible to identify the primary effects of Li or secondary actions due to the complexity
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of responses to Li toxicity. In conclusion, the exact mechanism behind the effects of Li

either as an inhibitor or stimulant of plant growth and physiology is still not clear;
however, a summary of reported and some suggested effects of Li are depicted, which
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show possible reasons for Li induced growth inhibition (Fig. 2).

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4. Li remediation strategies

Li is important from a research point of view especially due to its increasing

concentration in the environment and water. According to the Australian Capital
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Territory Environment Protection Regulation Li is listed as a pollutant that causes

environmental harm in irrigation water supplies (EPA, 2005). Its importance could not be
denied in studying scientific developments, plant nutrition as a trace element, and its
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clinical considerations in internal medicine (Freeman and Freeman, 2006; Kabata-

Pendias, 2010). Nonetheless, Li is being released into the environment in great amount
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due to its use in industries particularly as Li batteries (Al-Thyabat et al. 2013) or from
brines (Habashi 1997; Aral and Vecchio-Sadus, 2008). Disposal of Li batteries is a
challenging threat to the environment if it is not controlled and can be lethal to humans
and plants as well. The exposure of plants to Li toxicity evoked research on its potential
risks for cultivated lands, plants, animals, water resources, the environment, and
ultimately for human beings. The contamination of soil by Li is becoming a serious
problem and will be a threat for crop production in the near future. Additionally, lack of
considerable information about the tolerance mechanism of plants further amplifies the
 ACCEPTED MANUSCRIPT

situation. Therefore, there is a need to emphasize the research for prominent and
approachable solutions to minimize the entry of Li from its sources, especially from Li
batteries, into soil and the food chain. Li toxicity or its addition can be reduced in
different ways (Table 7). Possible remediation approaches can be employed with good
outcomes and so Li toxicity can be considerably managed.

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5. Conclusion
Studies are needed to know the exact role of Li in biological systems to elucidate
tolerance mechanisms because the effect of Li toxicity in many ecosystems is still poorly

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understood. The present article reviewed Li toxicity and related aspects including its
sources of contamination in soil and plants, its transportation from source to sink (food)
and its effects on plant growth, mineral nutrition and carbon assimilation. It can be

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summarized by saying that Li reduces plant growth by limiting plant biomass production,
and decreasing development of other morphological attributes. Various evidences also
highlight Li-induced alterations in numerous physiological functions and processes

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within plants such as alteration in antioxidant activity, higher production of ROS and
perturbation of the biological activity of important enzymes and, in severe cases,
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inhibition of DNA synthesis. Moreover, Li interferes with uptake and translocation of
some other biologically important elements. Tolerance mechanisms for Li are not yet
clear and a lot of research is required to elucidate tolerance mechanisms because the
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effects of Li in many ecosystems are still poorly described. Li pollution is becoming a

major environmental concern and has toxic effects on plants and animals including
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human beings. Remediation to this problem is imperative and needs further in-depth
study.
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Acknowledgements

The authors would like to thank Dr. Patrick Finnegan (Senior Lecturer, School of Plant
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Biology, University of Western Australia), for improving language and the worthy

reviewers for their helpful questions and suggestions which have made this paper better.
C
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References

Allender, W.J., Cresswell, G.C., Kaldor, J., Kennedy, I.R., 1997. Effect of lithium and
lanthanum on herbicide induced hormesis in hydroponically-grown cotton and
corn. J Plant Nutr. 20, 81–95.
 ACCEPTED MANUSCRIPT

Al-Thyabat, S., Nakamura, T., Shibata, E., Iizuka, A., 2013. Adaptation of minerals
processing operations for lithium-ion (LiBs) and nickel metal hydride (NiMH)
batteries recycling: critical review. Minerals Engineer, 45, 4-17.

Ammari, T.G., Al-Zu’bi, Y., Abu-Baker, S., Dababneh, B., Tahboub, A., 2011. The
occurrence of lithium in the environment of the Jordan Valley and its transfer into

PT
the food chain. Environmental geochemistry and health 33(5), 427-437.

Anderson, M.A., Bertsch, P.M., Miller, W.P., 1988. The distribution of lithium in

RI
selected soils and surface waters of the southeastern USA. Applied geochemistry
3(2), 205-212.

SC
Anjum, S.A., Ashraf, U., Khan, I., Tanveer, M. et al. 2016a. Chromium and aluminum
phyto-toxicity in maize; morpho-physiological responses and metal uptake.
Clean- Soil Air Water, DOI: 10.1002/clen.201500532

U
Anjum, S.A., Tanveer, M., Hussain, S., Bao, M., Wang, L., Khan, I., Ehsanullah, Samad,
AN
R.A., Tung, S.A., Shahzad, B., 2015a. Cadmium toxicity in maize (Zea mays L.):
Consequences on antioxidative systems, reactive oxygen species and cadmium
accumulation. Environmen- tal Science and Pollution Research, 22, 17022–
17030.
M

Anjum, S.A., Tanveer, M., Hussain, S., Ehsan ullah, Wang, L., Khan, I., Samad, R.A.,
Tung, S.A., Anum, M., Shahzad, B., 2015b. Morpho-physiological growth and
D

yield responses of two contrasting maize cultivars to cadmium exposure.

TE

CLEAN–Soil, Air. Water doi:10. 1002/clen.201400905.

Anjum, S.A., Tanveer, M., Hussain, S., Shahzad, B., Ashraf, U., Fahad, S., Hassan, W.,
Jan, S., Saleem, M.F., Khan, I., Bajwa, A.A., Wang, L., Mehmood, A., Samad,
EP

R.A., Tung, S.A., 2016b. Osmoregulation and antioxidant production in maize

under combined cadmium and arsenic stress. Environmental Science and
Pollution Research doi:10.1007/s11356-016-6382-1.
C

Aral, H., Vecchio-Sadus, A., 2008. Toxicity of lithium to humans and the environment—
AC

a literature review. Ecotoxicol. Environ. Saf. 70, 349–356.

Armbruster, A.M., Pullman, A., 1974. The effect of cation binding on the rotation barrier
of the peptide bond. FEBS Lett. 49, 18-21.


Bartolo, M.E., Carter, J.V., 1992. Lithium decreases cold-induced insoluble

depolymerization in mesophyll cells of spinach. Plant Physiol. 99, 1716–1718.
 ACCEPTED MANUSCRIPT

Bello, I., Haas, D., Bello, H.R., 1966. Interactions of protein-denaturing salts with model
amides. Biochemistry 5, 2539-2548.

Belyavskaya, N. A., 2001. Lithium-induced changes in gravicurvature,

statocyte ultrastructure and calcium balance of pea roots. In: Kiss,
J.Z., Kern, V.D. (eds.) Space life sciences: gravity perception and

PT
transduction in plants, fungi and unicellular organisms. Elsevier
Science Bv, Amsterdam, 961 966

RI
Berridge, M.J., 1993. Inositol triphosphate and calcium signaling. Nature 361, 315–325.

Bhattacharjee, S., 2005. Reactive oxygen species and oxidative burst: roles in stress,

SC
senescence and signal transduction in plants. Curr. Sci. 89, 1113–1121.

Bhattacharyya, B., Wolff, I., 1976. Stabilization of microtubules by lithium ion.

Biochem. Biophys. Res. Commun. 73, 383-390.

U
Bingham, F.T., Bradford, G.E., Page, A.L., 1964. Toxicity of lithium. Calif. Agric. 18,
AN
6–7.

Birch, N.J., (ed) 1991. Lithium and the cell: pharmacology and biochemistry. Academic
M

Press, San Diego

Birch, N.J., 1976. Possible mechanisms for biological action of lithium. Nature 204, 681.
D

Birch, N.J., 1988. Lithium, in: Seiler, H.G. (Ed.), Handbook on the Toxicity of Inorganic
TE

Compounds. Marcel Dekker, New York, pp. 382-393.

Birch-Hirschfeld, L., 1920. Untersuchungen uber die Ausbreitungsgeschwindiglfeit

geloster Stoffe in der Pflanze. yb. wiss. Bot. 59, 17.
EP

Bleiwas, D.I., Coffman, J.S., 1986. Lithium availability-market economy countries. In:
Circular, B.o.M.I. (Ed.), pp. 23.
C

Borovik-Romanova, T.F., 1965. The content of lithium in plants (Khitarov, N.I.,

AC

Jerusalem., 1969. Trans.). USSR: Inst. Geokem. Akad. Nauk.

Bowen, H.J.M., 1979. Environmental chemistry of the elements. New York: Academic
Press.

Bradford, V.G.R., 1963. Lithium survey of California's water resources. Soil Sci. 96, 77-
81.
 ACCEPTED MANUSCRIPT

Bradford, V.G.R., 1966. Lithium. In: CHAMPAN, H.D., (Ed.), Diagnostic Criteria for
Plants. Univ. of Cal. Div. of Agr. Sci. Chap. 17, pp. 218-224.

Brenchley, W.E., 1832. The action on the growth of crops on small percentages of certain
metallic compounds when applied with ordinary fertilizers. 1. Agric. Sci. 22, 704-
735.

PT
Cannon, H.L., Harms, T.F., Hamilton, J.C., 1975. Lithium in unconsolidated sediments
and plants of the Basin and Range Province, Southern California and Nevada (No.

RI
918). US Govt.

Carlier, G., Thellier, M., 1979. Lithium-perturbation of the induction of a methyl-glucose

SC
transport during aging of foliar disks of Pelargonium zonale (L.). Physiol. 7, 13-
26.

Chan, L.H., Sturchio, N.C., Katz, A., 1997. Lithium isotope study of the Yellowstone

U
hydrothermal system. EOS Trans. Am. Geophys. Union 78, F802 (abstr).
AN
Clemens, S., 2006. Evolution and function of phytochelatin synthases. J. Plant Physiol.
163(3), 319–332.
M

Codina, M.C., Morales, P.C., Serrano, G.M., 1983. Influencia de los elementos alcalinos
sobre el crecimiento y niveles de pigmentos cloroplásticos em plantas de Nicotina
rustica L. AD. Edafol. Agrobiol. 42, 601-612.
D

Dawson, E.B., 1991. The relationship of tap water and physiological levels of lithium to
TE

mental hospital admission and homicide in Texas. In Schrauzer, G.N.,

Klippel, K.F., (eds.): Lithium in Biology and Medicine. Weinheim: VCH
Verlag, pp 171 187.
EP

Dawson, E.P., Moore, T.D., McGanity, W.J., 1970. The mathematical relationship of
drinking water lithium and rainfall on mental hospital admission. Dis. Nerv. Syst.
C

31, 1–10.
AC

Dawson, E.P., Moore, T.D., McGanity, WJ., 1972. Relationship of lithium metabolism to
mental hospital admission and homicide. Dis. Nerv. Syst. 33, 546–556.

Desbiez, M.O., Thellier, M., 1977. Induced precedence between cotyledonary buds: Ionic
or ouabain treatments and memorization effects. In: Thellier, M., et al. (Ed.),
Transmembrane Ion Exchange in Plants. CNRS. Paris, pp. 607.

Desbiez, M.O., Thellier, M., 1978. Ionic control of the occurrence of a biological rhythm
for precedence between axillary buds. Physiol. 16, 785-798.
 ACCEPTED MANUSCRIPT

Dubey, R.S., 2005. Photosynthesis in plants under stressful conditions. In: Handbook of
Photosynthesis, Marcel Dekker, New York, pp. 859–875.

Duff, M.C., Kuhne, W.W., Halverson, N.V., Chang, C.S., Kitamura, E., Hawthorn, L.,
Stieve-Caldwell, E., 2014. mRNA Transcript abundance during plant growth and
the influence of Li+ exposure. Plant Science 229, 262-279.

PT
Edwards, I.K., 1941. Cytological studies of toxicity in meristem cells of roots of Zea
mays. II. The effects of lithium chloride. Proc. S.D. Acad. Sci. 21, 65-67.

RI
Efrati, S., Averbukh, M., Berman, S., Feldman, L., Dishy, V., Kachko, L., Weissgarten,
J., Golik, A., Averbukh, Z., 2005. N-Acetylcysteine ameliorates lithium-induced
renal failure in rats, Nephrol. Dial. Transplant. 20, 65–70.

SC
Enghag, P., (2008). Encyclopedia of the elements: technical data-history-processing-
applications. John Wiley and Sons.

U
Englemann, W., 1972. Lithium slows down the Kalanchoe clock. Z. Naturforsch. B:
AN
Anorg. Chem. Org. Chem. Biochem. Biophys. BioI. 27, 477-478.

Englemann, W., 1973. A slowing down of circadian rhythms by lithium ions. Z.

Naturforsch. C: Biochem. Biophys. BioI. Virol. 28, 733-736.
M

EPA. (2005). Australian capital territory environment protection regulation.

http:\\www.legislation.act.gov.au/sl/2005-
D

38/current/pdf/2005-38.pdf. Accessed April 15, 2010.

TE

Epstein, E., 1960. Calcium-lithium competition in absorption by plant roots. Nature 185,
705-706.
EP

Erakovic, V., Zupan, G., Varljen, J., Laginja, J., Simonic, A., 2000. Lithium plus
pilocarpine induced status epilepticus-biochemical changes, Neurosci.
Res. 36, 157– 166.
C

Eraslan, F., Inal, A., Savasturk, O., Gunes, A., 2007. Changes in antioxidative system and
AC

membrane damage of lettuce in response to salinity and boron toxicity. Sci. Hort.
114, 5-10.

Fouillac, C., Michard, G., 1981. Sodium/lithium ratio in water applied to geothermometry
of geothermal reservoirs. Geother. 10, 55-70.

Freeman, M.P., Freeman, S.A., 2006. Lithium: clinical considerations in internal

medicine. Am. J. Med. 119, 478-481.
 ACCEPTED MANUSCRIPT

Gaillochet, J., 1981. Effect of the lithium-chloride on the leaf

movements of Cassia fasciculata. Planta, 151, 544 548

Gumber, S.C., Loewus, M. W., Loewus, F.A., 1984. Further studies on myo-inositol-1-
phosphatase from the pollen of Lilium longiflorum Thunb. Plant Physiol. 76, 40-
44.

PT
Habashi, F., 1997. Handbook of extractive metallurgy. New York: Wiley-VCH

RI
Hallock, R.J., Hallock, L.L., (eds.) 1993. Detailed study of irrigation drainage in and near
wildlife areas, west-central Nevada, 1987-1990. Part B. effect on biota in Strill
water and Fernley Wildlife Service, US Bureau of Reclamation, and US Fish and

SC
Widelife Serivce, US Bureau of Reclamation, and US Bureau of Indian Affairs,
Carson City, NV.

U
Harpaz-Saad, S., Azoulay, T., Arazi, T., Ben-Yaakov, E., Mett, A., Shiboleth, Y. M., ...
& Eyal, Y. (2007). Chlorophyllase is a rate-limiting enzyme in chlorophyll
AN
catabolism and is posttranslationally regulated. The Plant Cell 19, 1007-1022.

Hassan, M.N., 1954. The effect of single salt solutions on the histogenesis of radish
M

seedlings. Alexandria J. Agric. Res. 2, 20-27.

Hawrylak-Nowak, B., Kalinowska, M., Szyma ńska, M., 2012. A study on selected
physiological parameters of plants grown under lithium supplementation. Biol.
D

Trace Elem. Res. 149, 425–430.

TE

Hem, J.D., 1985. Study and interpretation of the chemical

characteristics of natural water. USGS Water Supply Paper 2254. United States
Geological Survey, Alexandria, Virginia, USA
EP

Hem, J.D., 1992. Study and Interpretation of Chemical Characteristics of Natural Water
(3rd ed.), USGS Water-Supply Paper, pp. 2254.
C

Hu, J., Shi, G., Xu, Q., Wang, X., Yuan, Q., Du, K., 2007. Effects of Pb2+ on the active
AC

oxygen scavenging enzyme activities and ultrastructure in Potamogeton crispus

leaves. Russ. J. Plant Physiol. 54, 414–419.

Hull, S.L., Oty, U.V., Mayes, W.M., 2014. Rapid recovery of benthic invertebrates
downstream of hyperalkaline steel slag discharges. Hydrobiol. 736, 83-97.

Iqbal, K., Khan, A., Khan, K.M.M.A., 2004. Biological significance of ascorbic acid
(vitamin C) in human health—a review. Pak. J. Nutr. 3, 5–13.
 ACCEPTED MANUSCRIPT

Jacobson, L., Moore, D.P., Hannapel, R.J., 1960. Role of calcium in absorption on
monovalent cations. Plant Physiol., 35,352-357.

Jiang, L., Wang, L., Mu, S.Y., Tian, C.Y., 2014. Apocynum venetum: A newly found
lithium accumulator. Flora-Morphology, Distribution, Functional Ecology of
Plants, 209, 285-289.

PT
Johnson, E.N., (eds.)., 1975. Lithium research and therapy. New York: Academic Press,
pp. 569.

RI
Jones, A., Rogerson, M., Greenway, G., Potter, A.B., Mayes, W.M., 2013. Mine water
geochemistry and metal flux in a major historic Pb Zn F orefield, the Yorkshire

SC
Pennines, UK. Environ. Sci. Poll. Res.
20, 7570-7581

Jurkowska, H., Rogó ż, A., 1991. Uptake of lithium by plants as depending on soil

U
moisture content. Polish J. Soil Sci. 24, 93–97.
AN
Jurkowska, H., Rogó ż, A., Wojciechowicz, T., 1995. The effect of sodium on lithium
uptake by plants. Polish J. Soil Sci. 28, 135–138.
M

Jurkowska, H., Rogó ż, A., Wojciechowicz, T., 1998. Comparison of lithium toxic
influence on some cultivars of oats, maize and spinach. Acta Agraria et
Silvestria/Agraria 36, 37–42. (In Polish with English abstract)
D

Kabata-Pendias, A., 2010. Trace elements in soils and plants. CRC press.
TE

Kabata-Pendias, A., Mukherjee, A.B., 2007. Trace elements from soil to human.
Springer, Berlin, pp. 87–93.
EP

Kabata-Pendias, A., Pendias, H., 1992. Trace elements in soils and plants (2nd ed.). Boca
Raton, London: CRC Press.
C

Kalinowska, M., Hawrylak-Nowak, B., Szymańska, M., 2013. The Influence of two
lithium forms on the growth, L-ascorbic acid content and lithium accumulation in
AC

lettuce plants. Biological trace element research, 152(2), 251-257.

Kandeler, R., 1970. The effect of lithium and ADP on the phytochrome regulation of
flowering. Planta. 90, 203-207.

Kent, N.L., 1941. Absorption, translocation and ultimate fate of lithium in the wheat
plant. New Phytol. 40(4), 291–298.
 ACCEPTED MANUSCRIPT

Kergosien, Y., Thellier, M., Desbiez, M.O., 1979. Precedence between axillary buds in
Bidens pilosus L. Modeling at the macroscopic level in terms of catastrophes or at
the microscopic level in terms of a cellular "pump and leak". In: Delattre, P., et
al., (Ed.), Elaboration and Justification of Models. Paris: Malione, pp. 343.

Kielczykowska, M., Pastermak, K., Musik, I., Wroniska, J., 2004. The effect of lithium
administration in a diet on the chosen parameters of the antioxidant barrier in rats,

PT
Ann. Univ. Mariae Curie-Sklodowska 59, 140–145.

Klemfuss, H., Schrauzer, G.N., 1995. Effects of nutritional lithium deficiency on

RI
behavior in rats. Biol. Trace El. Res. 48, 131–139.

Kszos, L.A., Beauchamp, J.J., Stewart, A.J., 2003. Toxicity of lithium to three freshwater

SC
organisms and the antagonistic effect of sodium. Ecotoxicol. 12, 427-437.

Kszos, L.A., Stewart, A.J., 2003. Review of lithium in the aquatic environment:

U
distribution in the United States, toxicity and case example of groundwater
contamination. Ecotoxicol. 12, 439 447
AN
Laties, G.G., 1959. The development and control of coexisting respiratory systems in
slices of chicory root. Arch. Biochem. Biophys. 79, 378-391.
M

Lenntech, 2007. Lithium and water: reaction mechanisms, environmental impact

and health effects. /www.lenntech.com/elements-and-water/lithium-
D

andwater.htmS.
TE

Li, X., Gao, P., Gjetvaj, B., Westcott, N., Gruber, M.Y., 2009. Analysis of the
metabolome and transcriptome of Brassica carinata seedlings after lithium
chloride exposure. Plant Sci. 177, 68–80.
EP

Long, K.E., Brown, Jr, R.P., Woodburn, K.B., 1998. Lithium chloride: a flow-through
embryo-larval toxicity test with the fathead minnow, Pimephales promelas,
Rafinesque. Bull. Environ. Contam. Toxicol. 60, 312-317.
C

Magalhães, J.R., Wilox, G.E., Rocha, A.N.F., Silva, F.L.I.M., 1990. Research on lithium-
AC

phytological metabolism and recovery of hypo-lithium. Pesq. Agropec. Bras. 25,

1781–1787.

Makus, D.J., Zibilske, L., Lester, G., 2006. Effect of light intensity, soil type, and lithium
addition on spinach and mustard greens leaf constituents. Subtrop. Plant Sci. 58,
35–41.
 ACCEPTED MANUSCRIPT

Małecka, A., Piechalak, A., Morkunas, I., Tomaszewska, B., 2008. Accumulation of lead
in root cells of Pisum sativum. Acta Physiol. Plant 30(5), 629–637.

Mason, B., 1974. Principles of Geochemistry, 3rd ed. Wiley, New York

Mason, T.G., Maskell, E.J., 1931. Further studies on transpiration in the cotton plant. I.

PT
Preliminary observations on the transport of phosphorus, potassium and calcium.
Ann. Bot. Lond. 45, 125.

McStay, N.G., 1980. Effects of lithium on several plant systems. M.S. Thesis.

RI
Department of Botany, North Carolina State University, Raleigh, NC

Mikhaleva, L.M., Zhavoronkov, A.A., Kaktursky, L.V., Anke, M., Groppel, B., 2001.

SC
Oncomorphology in goats with acquired trace element deficiencies. In:
Biogeochemistry and Geochemical Ecology: Selected Presentations of the 2nd
Russian School of Thought: Geochemical Ecology and Biogeochemical Study of

U
Taxons of the Biosphere,” Moscow, January 25–29, 1999. Moscow: GUN NPC
AN
TMG MZ RF, pp. 190–197.

Millet, B, Badot, P., 1996. The revolving movement mechanism in

Phaseolus; New approaches to old questions. In: Greppin, H.,
M

Degli, Agosti, R., Bonzon, M., (eds.) Vistas on Biorhythmicity.

University of Geneva, Geneva, 77 98
D

Moore, S., 2007. Between rock and salt lake. Ind. Miner. June, 58 69.
TE

Mulkey, T.J., 2005. Alteration of growth and gravitropic response of maize roots by
lithium. Grav. Space Biol. 18(2), 119–120.
EP

Naranjo, A., Romero, C., Bellés, J.M., Montesinos, C., Vicente, O., Serrano, R., 2003.
Lithium treatment induces a hypersensitive-like response in tobacco. Planta 217,
C

417–424.
AC

Nciri, R., Allagui, M.S., Bourogaa, E., Saoudi, M., Murat, J.C., Croute, F., Elfeki, A.,
2012. Lipid peroxidation, antioxidant activities and stress protein (HSP72/73,
GRP94) expression in kidney and liver of rats under lithium treatment. J. Physiol.
Biochem. 68(1), 11-18.

Oktem, F., Ozguner, F., Sulak, O., Olgar, S., Akturk, O., Yilmaz, H.R., Altuntas, I., 2005.
Lithium-induced renal toxicity in rats: protection by a novel antioxidant caffeic
acid phenethyl ester, Mol. Cell. Biochem. 277, 109–115.
 ACCEPTED MANUSCRIPT

Ono, T., Wada, O., 1991. Effect of lithium deficient diet on avoidance behavior of
animals and a discussion of the essentiality of lithium. Biomed. Res. Trace El. 2,
264–265.

Pickett, E.E., 1983. Evidence for the essentiality of lithium in the rat. In: Anke, M., et al.,
1983. (Ed.): “Proceedings 4. Spurenelement Symposium.” Jena: VEB

PT
Kongressdruck, pp 66–70.

Powell, I.T., Richards, E.G., 1972. Specific effects of lithium on stacking equilibria in

RI
polynucleotides. Acta Biochim. Biophys. Acad. Sci. Hung. 281, 145-151.

Ribas, B., 1991. Lithium, in: E. Merian (Ed.), Metals and their Compounds in the

SC
Environment, 4th edn., VCH, Weinheim, pp. 1014-1023.

Richard, M.J., Belleville, F., Chalas, J., Ceballos-Picot, I., Vitoux, D., Boyer, M.J.,
Haudiere, J., Favier, A., 1997. Glutathione peroxidases: value of their

U
determination in clinical biology, Ann. Biol. Clin. 55, 195– 207.
AN
Robinson, W.O., Steinkoenig, L.A., Miller, C.F., 1971. The relation of some of the rarer
elements in soils and plants. U.S. Dept. Agric. Bull. No. 600 (p. 27). Washington,
DC: U.S. Government Printing Office.
M

Rogo´z, A., 2004. Studies on lithium forms in soil. Part II. Relationship between forms in
soil and their assimilativeness to plants. J. Elementology 9, 727-734.
D

Rusin, G.G., 1979. Application of lithium salts for increasing yield and quality of
TE

hydroponically grown cucumbers. Nauchnye Trudy Ukr. S. Kh. Akad. 228, 58-
61.
EP

Schrauzer, G.N., 2002. Lithium: Occurrence, Dietary Intakes, Nutritional Essentiality. J.

Am. College Nut. 21, 14-21.

Shacklette, H.T., Boerngen, J.G., 1984. Element Concentration in Soils and Other
C

Surficial Materials of the Conterminous United States,” Geological Survey

AC

Professional Paper (United States), pp. 1270.

Shacklette, H.T., Erdman, J.A., Harms, T.F., Papp, C.S.E., 1978. Trace elements in plant
foodstuffs. In: Oehme, F.W., (Ed.), Toxicity of Heavy Metals in the Environment,
4th Edn., Marcel Dekker, New York, pp. 25-68.

Shkolnik, M.Y.A., 1984. Trace Elements in Plants, Amsterdam, Elsevier pp. 463.
 ACCEPTED MANUSCRIPT

Singh, R., Tripathi, R.D., Dwivedi, S., Kumar, A., Trivedi, PK., Chakrabarty, D., 2010.
Lead bioaccumulation potential of an aquatic macrophyte Najas indica are related
to antioxidant system. Bioresour. Technol. 101, 3025–3032.

Siroonian, H.A., Shaw, D.M., Jones, R.E., 1959. Lithium geochemistry and the source of
spodumene pegmatites of the Preissae-Lamotte-Lacorne region ofwestern

PT
Quebec. Can. Minereralogist 6, 320-328.

Sposito, G., 1986. Distribution of potentially hazardous trace metals. In: Sigel, H., (Ed.),

RI
Metal Ions in Biological Systems, Vol. 20, 4th edn., Marcel Dekker, New York,
pp. 1-20.

SC
Stevenson, J.M., Perera, I.Y., Heilmann, I., Persson, S., Boss, W.F., 2000.
Inositol signaling and plant growth. Trends Plant Sci. 5,252 258

U
Stewart, A. J., Kszos, L. A., 1996. Caution on using lithium (Li+) as a conservative tracer
in hydrological studies. Limnol. Oceano. 41, 190-191.
AN
Stracher, A., 1960. Deuterium exchanges or ribonuclease and oxidized ribonuclease in
strong salt solutions. C.R. Trav. Lab. Carlsberg, 30, 468-481.
M

Tandon, A., Dhawan, D.K., Nagpaul, J.P., 1998. Effect of lithium on hepatic lipid
peroxidation and antioxidative enzymes under different dietary protein regimens.
D

J. Appl. Toxicol. 18, 187– 190.

TE

Tate, C.M., Broshears, R.E., McKnight, D.M., 1995. Phosphate dynamics in an acidic
mountain stream: interactions involving algal uptake, sorption by iron oxide, and
photoreduction. Limnol. Oceano. 40, 938-946.
EP

Thellier, M., Desbiez, M.O., 1977. Model of a switching "on" and "off' pump and leak, as
a relay and amplification mechanism in the control of morphogenesis. In: E.
Marre, E., et al., (Eds.), Regulation of cell membrane activities in plants.
C

Amsterdam: Elsevier. North-Holland Biomedical Press, pp. 332.

AC

Thellier, M., Thoiron, B., Thoiron, A., Le Guiel, J., Luttge, U., 1980. Effects of lithium
and potassium on recovery of solute uptake capacity of Acer pseudoplata- nus
cells after gas shock. Physiol. Plant. 49, 93-99.

To¨lgyesi, G., 1983. Distribution of lithium in Hungarian soils and plants. In: Anke, M.,
et al., (Eds.), Lithium 4. Spurenelementsymposium. Jena: Friedrich Schiller
Universita¨t, pp. 39–44.
 ACCEPTED MANUSCRIPT

United States. Department of Energy, Jeppson, D.W., 1978. Lithium literature review:
lithium's properties and interactions (p. 26). Department of Energy.

Vergara-edwards, L., Parada-fre- Derick, N., Pavlovick-zuvic, P., 1986.

Recovery of lithium from cristallized salts in the solar evaporation of solar in
Atacama brines. Lithium Curr. Appl. Sci. Me. Teehnol. 53, 47-59.

PT
Vine, J.D., Dooley, J.R. Jr. 1980. Where on Earth is all the lithium?, with a section on
Uranium isotope studies. Fish Lake Valley, Nevada,U.S. Geological Survey

RI
Open-File Report 80-1234

Vlasyuk, P.A., Kuz'menko, L.M., 1975. Metabolic activity of potato plant ribo- somes in

SC
dependence on their supply with lithium. Fiziol. Biokhim. Kun. Rast. 7, 563-568.

Vlasyuk, P.A., Kuz'menko, L.M., Okhrimenko, M.E., 1975a. Content and fractional
composition of potato protein and nucleic acids under lithium effect. Dopov.

U
Akad. Nauk. Ukr. RSR. Ser. B: Geol. Geofi,z. Khim. Bioi. pp. 742-748.
AN
Vlasyuk, P.A., Kuz'menko, L.M., Okhrimenko, M.F., 1979. The role of lithium in
protein-nucleic acid metabolism in plants. Fiziol. Biokhim. Kul't. Rast. 11, 438-
447.
M

Vlasyuk, P.A., Okhrimenko, M.E., Kuz'menko, L.M., 1975b. Fractional and amino acidic
compositions of proteins and content of free amino acids in potato under the
influence of lithium. Fiziol. Biokhim. Kul't. Rast. 7, 115-120.
D

Vlasyuk, P.A., Okhrimenko, M.F., Kuzmenko, L.M., 1973. Effect of lithium on contents
TE

and composition of organic acids in plants of the Solanaceae. Fiziologiya

Biokhimiya Rastenii 5,121-124. (English summary).
EP

Weber, U., Engelmann, W., Mayer, W.E., 1992. Effects of tetraethylammoniumchloride

(TEA), vanadate, and alkali ions on the lateral
leaflet movement rhythm of Desmodium motorium (Houtt) Merr.
C

Chronobiol. Int. 9,269 277

AC

Weeks, M.E., 1956. Discovery of the elements (6th ed.). J. Chem. Educ. Easton. pp. 578.

Williams, R.J.T., 1973. The chemistry and biochemistry of lithium. In: Geershon, S., et
al., (Ed.), Lithium. Its role in psychiatric research and treatment. New York:
Plenum Press, pp. 358.

Wortley, W.R.S., 1936. Report of research, 1934-36. The effect of salts of lithium on the
resistance of certain plants to disease. l.R. Agric. Soc. Engl. 97, 492- 498.
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Zachariassen, E., Johnsson, A., 1988. Effects of lithum ions on the

circumnutations of Helianthus hypocotyls. Plant Physiol.
72, 147 152

Zeller, S., Fuller, U., 2000. Long-distance transport of alkali metals in maturing wheat.
Biol. Plant 43(4), 523–528.

PT
Zonia, L.E., Tupy, J., 1995. Lithium treatment of Nicotiana tabacum
microspores blocks polar nuclear migration, disrupts the partitioning of

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membrane-associated Ca2+ , and induces symmetrical mitosis. Sex Plant Reprod.
8,152 160

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Table 1: Different identified resources of Li entry in ecosystem

Li source Li concentration Reference
Li Brine 20 mg l-1 in dead sea Habashi 1997; Moore 2007
1500 mg l-1 in Salar de Atacama

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Li pegmatite and/ Li brines 7.6 ×106 kg Cannon et al., 1975; Evans, 1987

Authigenic clays 200 – 500 ppm in detrital clays Chan et al., 1997

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Li shaley rocks 20-100 mg kg-1 Vine and Dooley 1980

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Land disposal of Li-hydride and Li-deuteride Kszoz and Stewart 2003
material

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Effluents from fabrics, ceramics and Long et al., 1998; Enghag 2008

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cosmetics industry

Table 2: Li concentration in different water bodies

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Li concentration Water body Location of water sample Reference
1-10 µg l-1 Surface water - Mason 1974; Lenntech 2007

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0.17 mg l-1 Sea water - Mason 1974; Lenntech 2007
Kszos et al., 2003

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0.5 mg l-1 Ground water - Mason 1974; Lenntech 2007
<0.04 mg l-1 Surface water - Kszos et al., 2003

500 g l-1
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Ground water Northern Chile Saunders 1985; Schrauzer 2002
0.07 – 40 µg l-1 Fresh water - Bowen 1979; Sposito 1986
Upto 100 mg l-1 Natural mineral waters Texas, USA Dawson 1991
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≤0.05 mg l-1 Well water and few surface California, USA Bradford 1963
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water samples
10-19 mg l-1 Ground water USA Kszos and Stewart 2003
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Table 3: Li concentrations in different soil types

Experimental location Soil type/Soil series Li concentration Reference
(mg/kg)/(mg/L)
USA Cecil (Clayey,kaolinitic,thermic 11.49 Anderson et al. 1988
TypicHapludult)

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USA Iredell (Fine, mont morillonitic, 25.38
thermic Typic Hapludalf)

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USA Madison (Clayey, kaolinitic, thermic 11.93
Typic Hapludult)

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USA Louisa (Fine, mont morillonitic, 33.29
thermic Ruptic-Ultic

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Dystrochrept)

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USA Wakulla (Sandy,siliceous,thermic 5.82
Psammentic Hapludult)

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USA Bonifay (Loamy, siliceous, thermic 3.74
Grossarenic Plinthic Paleudult)
Jorden valley Ustochreptic and ustollic camborthids 1.04-1.56 Ammari et al. 2011

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and calciorthids, ustic torriorthents.

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Northern Jorden Valley Ustochreptic and ustollic camborthids 1.05-1.06
and calciorthids, ustic torriorthents.
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Middle Jorden Valley Ustochreptic and ustollic camborthids 1.19-2.68
and calciorthids, ustic torriorthents.
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Southern Jorden Valley Ustochreptic and ustollic camborthids 0.95-1.48

and calciorthids, ustic torriorthents.
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Information in parenthesis is explaining soil family classification of different soils studied in USA (Anderson et al. 1988)
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Table 4: Li concentration in different food crops and daily intake

Crop plant Li concentration/mg kg-1 Estimated daily intake (mg Li/day) References

Lettuce 0.3-0.6 (FW) 0.09-0.18 Amari et al. 2011

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Cabbage 1.2 (FW) 0.12-0.48
Green onion 1.8 (FW) 0.18
Spinach 4.6 (FW) 1.15-1.38

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Forage plant species 0.2-200 (FW) - To lgyesi, 1983
Fodder beat 0.3-11.7 (FW) - Rogo’z,2004

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Celery 6.6 (DM) - Kabata-Pedias, 1992
Chard 6.2 (DM) - Kabata-Pedias, 1992

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Table 5: Response of different plant species under low and high lithium toxic level
Family Plant species Toxicity level Effects Reference

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Apocynaceae Apocynum No reduction in A. venetum shoot and root dry weight, chlorophyll contents and Jiang et al.
Low (50 mg kg-1)

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venetum leaf gas exchange 2014
High (200 and 400 mg Significant reduction in shoot and root dry weight, chlorophyll contents and leaf
kg-1) gas exchange

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Asteraceae Helianthus Low (20 and 40 mM) No reduction in hypocotyl length and in circumnutation Stolarz et al.

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annuus Hypocotyl length was reduced by 34 and 55% respectively and circumnutation 2015
(Sunflower) High (60 and 80 mM)
was reduced by 30 and 70% respectively.
Asteraceae Lactuca sativa Low (2.5 mg dm-3) Significant increase in root system Kalinowska et
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var. capitata al. 2013
High (50 or 100 dm-3) Considerable reduction in root system
(Lettuce)
Poaceae Zea mays Low (5 mg dm-3) Shoot biomass was increased by 15% Hawrylak-
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(Maize) High (50 mg dm-3) Shoot biomass was reduced by 32% Nowak et al.
Low (5 mg dm-3) 2012
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Asteraceae Helianthus Shoot biomass was increased by 10%

annuus
High (50 mg dm-3) Shoot biomass was reduced by 27%
(Sunflower)
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Table 6: Influence of different Li concentrations on plant growth reduction (Bingham et al. 1964)
Plant species Li concentration for normal Li concentration at which Li concentration at which
growth (ppm) 25% growth reduced (ppm) growth completely reduced
(ppm)

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Avocado (very sensitive) <5 6 100-450
Soybean (very sensitive) <5 7 100-300

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Sour orange (very sensitive) <5 8 100-300
Grape (sensitive) <5 12 300-600

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Red kidney bean (sensitive) <5 12 100-400
Cotton (slightly sensitive) <5 25 500-2000
Dallis grass (slightly sensitive) <5 25 300-500

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Red beat (slightly sensitive) <10 35 2500-4000
Rhodes grass (tolerant) <10 65 2500-3000

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Sweet corn (tolerant) <5 70 350-500

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Table 7: Proposed remediation strategies and their outcomes

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Remediation strategy Outcomes

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1) Identification and implementation of Li hyper-accumulator species to Outcomes would be identification of some fungi like Aspergillus or
sequester Li Arbuscular mycorrhizial fungi can accumulate high concentrations of Li and

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it contributes considerably in remediation of Li toxicity in soils. These living
organisms can produce some acids in soil that can act as chelating
compounds.

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2) Soil amendment practices such as organic matter addition to soils This could be another strategy to reduce Li toxicity in soil because organic
matter has ability to bind Li. Further research would define the interaction of

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Li reactivity and organic matter.

3) Application of chelators or chaperones This would highlight the potential of chelators or chaperones to ameliorate
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Li toxicity.

4) Screening and identification of Li tolerant genotypes This would aids in understanding the molecular basis of Li tolerance in those
Li stress tolerant genotypes.
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5) Identification and use of Li accumulator plants This would highlight the potential of the use of these accumulators and give
better insight into the Li uptake, translocation, distribution and detoxification
by these plant species (e.g. Apocynum venetum- Jiang et al. 2014)

6) Ex-situ and in-situ immobilization techniques for the remediation of Li- Outcome would indicate the potential of these techniques for Li remediation
contaminated soils. however storage and disposal of scraped Li-contaminated soil and high
invasively to the environment could be major problems for ex-situ soil
treatment.
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7) Soil stabilization and solidification Results would suggest that the addition of some regents and binding

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substances to a contaminated material to communicate dimensional/physical
stability to contain contaminants in a solid product and decrease access by
exterior substance through contamination of chemical reaction, reduce
permeability and encapsulation

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8) Removal of excess Li from ground water can be done using resins This would highlight and identify the substances (e.g. resins-Kszos and

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Stewart, 2003) to remove the Li from Li contaminated water.

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Figure 1: This figure is representing results of Li toxicity; fig (A) is showing the development of necrotic spots on the older leaves of lettuce plant when exposed

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to Li concentration of 100 mg Li dm-3 (Kalinowska et al. 2013); fig (B) is showing the comparison of the development of necrotic spot on sunflower seedling as
compared to control (Hawrylak-Nowak et al. 2012); fig (C) is illustrating the toxic effect of chlorophyll a, b, and carotenoid contents in sunflower and maize and
showed that with the increase in Li concentration significant reduction in these photosynthetic pigments were observed (Hawrylak-Nowak et al. 2012).

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Fig 2: Possible mechanism in which Li reduced the plant growth by disrupting numerous plant physiological processes. Li disturbs the balance between
production of reactive oxygen species (ROS) and antioxidant defense causing accumulation of reactive oxygen species, which induces oxidative stress. Higher
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ROS production also influences enzymatic activity. Li on other hand alters the translation and transcription process which reduces biosynthesis of amino acids,
lipids, and DNA. Increase in Li concentration in the vicinity of roots results in inhibited root growth due to higher ethylene production and development of
necrotic spots. In shoot and leaves, higher Li concentration activates chlorophyllase enzymes and disrupts chlorophyll contents, results in early leaf abscission,
which reduces the carboxylation and leaf gas exchange.
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Highlights

Lithium might be a serious hazardous pollutant for soil and environment in future.

Lithium stimulates or reduced plant growth depending on its concentration.

Bioavailability of Lithium depends on numerous factors.

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Lithium disrupts numerous metabolic processes such as photosynthesis, DNA biosynthesis and
enzyme activation in plants.

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B. Shahzad and M. Tanveer collected, compiled and wrote information, W. Hassan, A. Shah
gave useful suggestion and improved the text, S.A. Anjum, S.A. Cheema and I. Ali improved
grammatical and also discussed important information in manuscript.

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