Plant Mineral Nutrition: July 2014
Plant Mineral Nutrition: July 2014
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. Acknowledgements
Several inorganic minerals are essential for plant growth by chemical analysis of the previously digested leaf or
and these are usually obtained by roots from the soil. whole plants. Specific parts of the plant, like seeds, accu-
Availability of minerals in the soil is determined by the mulate some microelements to much higher concentrations
physical and chemical characteristics of the soil. Plants – for example, the zinc or iron levels can be 10–20 times
higher than the values given in Table 1. See also: Plant
can directly influence nutrient availability around the
Macro- and Micronutrient Minerals
root surface; this zone is called the rhizosphere. Plants
The elements that are essential for growth serve both
adjust root architecture and exudation according to their structural and biochemical roles in the plant and many
nutrient requirements and under deficiency these chan- have multiple functions. It is difficult to specifically classify
ges can be a marker for nutrient status. Nutrients are the role of each nutrient, but they can be placed into general
taken up from the soil using plasma-membrane located types based on function (see Table 1). Type I nutrients are
transporter proteins and excess is stored in the cell bound into the structure of carbon compounds, such as
vacuole or converted into polymerised storage forms. For nucleic acids and proteins. Type II nutrients are required
crops it is essential to match nutrient supply to demand for energy storage and transport. Type III nutrients are
throughout the growth season to obtain the maximum linked with cell wall structure and Type IV are integral as
yield. These nutrient storage forms can be used as agri- constituents of enzymes or other molecules required for
metabolism (e.g. chlorophyll and ferredoxin). Type V can
cultural indicators of crop nutrient status and the
activate enzymes or control their activity and Type VI
potential for fertilizer leaching losses. Membrane trans-
nutrients serve as major cellular osmotica.
porters provide a gateway for nutrient entry into plants,
but the selectivity of these filters can breakdown when
chemically similar minerals are present at very high con- Nutrient Availability
centrations. The minerals may not be essential for The physical and chemical characteristics of the soil
growth, but they can enter plant cells and cause toxicity. determine the availability of nutrients to uptake by plant
roots. The nutrients dissolved in soil water are those that
are generally available for uptake and this is why most
plants can be successfully grown in hydroponics or water
Introduction culture. In soil the plant can directly influence nutrient
availability in the area around the root surface; this zone is
Plants require some specific elements from the external
called the rhizosphere. Root-mediated localised changes in
environment and these are usually obtained by roots from
pH and soil microbes can directly influence the water
the soil. Less typically nutrients can also be taken up across
solubility of many nutrients. For example, a more acidic
the surface of leaves or in specialised structures in the few
rhizosphere pH dissolves soil mineral phosphorus,
types of plants that can catch and digest insects. The
increasing solubility and making the nutrient available for
quantities of each element required by plants can be used to
uptake by plant roots. A plant can adjust these root
define them as being either macro- or micro-nutrients. A
properties according to their nutrient requirements and
complete list of the mineral nutrients found in plants is
under deficiency these changes can be a marker for nutrient
shown in Table 1, and this type of information is obtained
status. Many plants (e.g. oilseed rape – Brassica napus)
excrete organic acids under deficiency (e.g. phosphorus
eLS subject area: Plant Science and iron) to increase the soil availability of these nutrients.
Furthermore, some plant roots excrete specific enzymes
How to cite: and chelating molecules to improve soil nutrient avail-
Miller, Anthony J (July 2014) Plant Mineral Nutrition. In: eLS.
ability, for example, phosphatases for phosphate and
John Wiley & Sons, Ltd: Chichester.
DOI: 10.1002/9780470015902.a0023717
siderophores for iron. Roots can encourage the growth of
particular types of bacteria and fungi that can solubilise
branch locally increasing the area for uptake and short- Transport; Plant–Fungal Interactions in Mycorrhizas;
ening the distance for diffusion (Mommer et al., 2012). The Root Nodules (Legume–Rhizobium Symbiosis)
upper layers of soil usually contain the highest amounts of
nutrients and so the top soil is frequently densely covered
with roots. Phosphorus is an example of a nutrient that
occurs in the top soil and bean roots proliferate in this layer
Homoeostasis, Storage and
to acquire this nutrient (Lynch and Brown, 2001). Primary Measuring Nutrient Status
roots grow deeper through the profile to exploit the water
and nutrient (e.g. nitrate) supplies that are often located The interplay between plasma membrane uptake and
much lower in the soil. See also: Roots and Root Systems nutrient storage forms is balanced to maintain cytoplasmic
Plants have gene families of nutrient transporters that concentrations that are optimised for plant growth (Figure
each specialise in the uptake of soil available nutrients, for 2). Nutrient homoeostasis is the regulation of supply to
example, ammonium and nitrate. There are key steps in the maintain an optimum environment for cellular biochem-
uptake of nutrients, these are transport across the plasma ical reactions in the cytoplasm. The balance or homo-
membrane from the soil into root cells, storage in the eostasis is important for a plant, yet very little is known
vacuole and loading of the long distance transport systems about how this cellular equilibrium is sensed and regulated
in plants (phloem and xylem), unloading into the growing within individual plant cells. It has been proposed that
tissues such as leaves or seeds. Each of these transport steps changes in root cytoplasmic nutrient concentrations can
is mediated by specific transporter proteins and they are signal nutrient status (e.g. nitrate, Miller and Smith, 2008;
potential regulatory steps in the pathway for nutrient entry potassium, Walker et al., 1996).
into the plant. This regulation can occur by changes in gene Excess nutrients are usually stored in cell vacuoles and
expression or by post-translational regulation of the pro- elemental analysis of whole tissue can be used to measure
tein. Often mutations in the transporter results in plants nutrient pools. For example, manganese (Mn) tissue
that demonstrate nutrient deficiency symptoms under accumulations can vary between plant species growing on
normal supply conditions that would be adequate for a the same soil. The general critical threshold concentration
wild type plant. But sometimes the mutation is hidden for the onset of Mn deficiency in leaves is in the range of 10–
because another transport in the family can step up 20 mg kg21 and it can particularly be a problem for plants
expression to compensate for the missing function. Some of to acquire the metal in alkaline soils (Marschner, 1995). At
the plasma membrane transporters are also involved in the other extreme, some rare plants can hyper-accumulate
sensing the external availability of nutrients and these have leaf manganese to 10 000 mg kg21, for example, Gossia
been described as ‘transceptors’ (Gojon et al., 2011). For a species (Fernando et al., 2009). Nutrients can sometimes be
recent list of the plant nutrient transporter families check stored after conversion into specific forms, and the
the ARAMEMNON website http://aramemnon.botani- amounts of these storage forms can also be used as indi-
k.uni-koeln.de/. Although the gene families of nutrient cators of a plant’s nutrient status. For example, phos-
transporters have selectivity for each type of nutrient this phorus can be stored as phytate in seeds, but the nutrient
specificity can break down when concentrations of other can be stored in all cells to a lesser extent as phosphate in
ions become high enough (see Section ‘Deficiency and the vacuole. Iron storage has been linked to accumulation
Toxicity’ below). The selectivity and transport rate of a of the protein ferritin, particularly in legume seeds like peas
plasma membrane nutrient transporter can also be and beans, but this may result from iron toxicity rather
important for determining crop nutrient use efficiency. than a direct role in storage of the metal (Briat et al., 2010).
Some nutrients are acquired by beneficial interactions or Nutrient transporters at the vacuolar membrane mediate
symbioses with bacteria and/or fungi. In symbiosis with the storage and remobilization of nutrients, and like at
fungi, the plant benefits from the increased surface area the plasma membrane, there are specific families of pro-
provided for mining nutrients from the soil, this has teins mediating these steps (Martinoia et al., 2012). The
obvious advantages for the acquisition of soil immobile activity of these vacuolar transporters must be regulated in
minerals, like phosphorus, but also for water uptake. concert with those at the plasma membrane to achieve
Together with bacteria, legumes can form a nitrogen-fixing homoeostasis and optimal cytoplasmic concentrations of
nodule that supplies nitrogen to the plant taken from the nutrients for metabolism and growth. Very high con-
air. A range of these symbiotic associations occur and their centrations of some nutrients can result in toxicity in plants
importance to each individual plant for nutrient acquisi- (see Figure 2) and this occurs when the storage capacity is
tion depends on the local availability of nutrients. In some exceeded by the supply. See also: Heavy Metal Adaptation;
nutrient-poor soils a symbiotic relationship may be essen- Iron in Plants
tial for the plant to grow and reproduce. These relation- The nutrient status of plants can be identified by whole
ships can also be important for plants growing in toxic tissue digests followed by elemental analysis. Young leaf
environments, for example, there are some plants that material is usually used for this analysis because this tissue
tolerate soils containing heavy metals by forming fungal is a good indicator of the whole plant nutrient status. The
mycorrhizal associations. See also: Improving Nutrient concentration in dry matter of a given nutrient can then be
Use Efficiency in Crops; Plant Nitrogen Nutrition and compared with tabulated values of the optimal range for
40 Deficiency with
symptoms
each type of crop. As this laboratory analysis can be time- more specifically which nutrient is missing. For example,
consuming, simpler assays have been developed that can be nitrogen deficiency results in a general chlorosis, but iron-
used by farmers in a field. Small deficiencies in nutrients deficient plants showing yellowing in the leaf between
may not present obvious symptoms, but can result in a sub- veins. When specific elements, like phosphorus, are defi-
maximal crop yield for farmers. The general nutrient status cient plants will develop a purple colouration due to the
of a crop can be assessed by the ‘greenness’ or chlorophyll production of large amounts of anthocyanins (see Figure 3,
content using hand-held or tractor mounted devices for right). These chemicals are produced when plants are
measuring this parameter. Nitrogen supply is well-known stressed and they have a biochemically protective role in
to influence the greenness of a crop. Insufficient phos- the cell. Tissue death or necrosis follows chlorosis as defi-
phorus supply can also influence on wheat leaf chlorophyll ciencies become more acute. In potassium-deficient plants
content and grain yield (Figure 3, left). The down-side of this necrosis occurs along leaf margins, but in manganese-
chlorophyll assessment method is that other factors like deficient plants necrosis occurs between veins. For some
pathogens or insect attack can complicate the measure- nutrients such as iron, the deficiency symptoms show first
ment. Plant leaf or petiole sap analysis can be used as an in a young leaf; this suggests that the element is not easily
indicator of nutrient status and portable equipment to translocated from old to young leaves. Nitrogen, potas-
measure nutrient concentrations in the field is available. sium and magnesium are easily loaded into the phloem and
Like whole tissue digests, sap analysis results can be com- xylem and translocated from old leaves to younger devel-
pared with previously determined optimal nutrient con- oping leaves. For these nutrients, the older leaves exhibit
centrations to decide if the crop needs more fertilizer. the deficiency symptoms. See also: Anthocyanins; Plant
Macro- and Micronutrient Minerals
As described above, very high concentrations of some
Deficiency and Toxicity nutrients can result in plant toxicity (see Figure 2) although
this rarely occurs in nature. The normal entry route for
As nutrients concentrations are critical for the processes nutrients can become a way for toxic elements to enter
involved in metabolism and growth, a limited supply of any plants. For example, transporters for the uptake of trace
nutrient will result in suboptimal stature and yield. element metals can become a pathway for the entry of other
Nutrient deficiencies result in stunted growth, but there are toxic metals like cadmium and under saline conditions
often other visual symptoms. Deficiency symptoms com- sodium can enter through potassium transporters as the
monly include chlorosis, a yellowing of the leaf and stem external abundance of sodium compared to potassium is so
tissues. The precise pattern of the chlorosis can indicate high. The accumulation of these toxic elements in plants
2
1 Low P
0
0 10 20 30 40 50 60 70 80 Strawberry leaf
Days after anthesis with sufficient P
P nutrition affects wheat grain yield
80
grain dry weight (mg)
Central spikelet
60 Sufficient P
40
Low P
20
0
0 10 20 30 40 50 60 70 80
Strawberry leaf
Days after anthesis showing P deficiency
Figure 3 The influence of varying phosphorus supply on wheat leaves and yield (left) and visual leaf symptoms in strawberry (right). Wheat figure shows
the effects of phosphorus supply on chlorophyll (upper) and grain yield (lower). Reproduced with permission from Kochian L (2002) Molecular physiology
of mineral nutrient acquisition, transport, and utilization. In: Bob B, Wilhelm G and Russell J (eds.) Biochemistry & Molecular Biology of Plants. Oxford,
Oxfordshire: John Wiley & Sons. ISBN 978-0-943088-39-6. & John Wiley & Sons.
can be ameliorated by increasing the supply of the nutrient Root strategies for improving nutrient acquisition
that usually enters the plant via this transporter system.
See also: Potassium in Plants • Rhizosphere - exudates,
acidification, enzymes and
specific types of microbes.
Future Prospects for Crop Nutrition • Uptake - transporters and
regulation.
When a nutrient deficiency can be detected in a crop there is • Root architecture - root hairs,
laterals, surface, depth and
already likely to be a yield penalty. In other words, nutrient penetration.
status detection in plants is too late; therefore, new meth- • Symbioses - mycorrhization and
ods for measuring and predicting soil nutrient availability nodulation.
are important for the early diagnosis of impending nutrient
limitations. This approach can ensure that the soil avail-
able nutrients are maintained at optimal levels for the crop, Figure 4 Some future strategies for improving crop nutrient acquisition.
but not at levels that result in fertilizer leaching. The
nutrient status of soil is usually assessed by taking core
samples in the field, then measuring the extracted nutrients availability. Choosing the appropriate type of bacteria can
in the laboratory. The accuracy of this method depends on encourage root growth and health, promoting nutrient
the type of extraction used, but often it overestimates the cycling in the rhizosphere. Soil and seed could be inocu-
amount of the nutrient that is actually available for uptake lated using beneficial microbes, with the species mix tai-
by roots. New more accurate methods for measuring soil lored for each type of crop. In addition, the ability of the
nutrient availability are important for improving crop plant to host particular types of bacteria may be enhanced
nutrition. by the root exudate composition. There is some evidence
Figure 4 shows some of the future possible strategies for that modern crop cultivars may have lost some of the
improving nutrient use by crop roots. For example, the chemical exudates found in ancestral varieties that can
inoculation of soil with specific types of bacteria tailored influence nutrient availability in the rhizosphere. For
for each crop may be a way of enhancing the nutrient example, root exudates can contain nitrification inhibitors
blocking the conversion of ammonium into nitrate (Sub- Hardoim PR, Andreote FD, Reinhold-Hurek B et al. (2011) Rice
barao et al., 2009). The plasma membrane transporters root-associated bacteria: insights into community structures
that mediate the uptake of nutrients by roots may also be across 10 cultivars. FEMS Microbiology Ecology 77: 154–164.
targets for improved acquisition of nutrients. This may be Hirsch PR, Mauchline TH and Clark IM (2010) Culture-inde-
achieved by increasing the affinity of the transporter for the pendent molecular techniques for soil microbial ecology. Soil
nutrient or by modifying the regulation of transporter Biology and Biochemistry 42: 878–887.
activity, for example, by phosphorylation state of the Lynch JP and Brown KM (2001) Topsoil foraging – An archi-
protein. Root architecture is another potential target as tectural adaptation of plants to low phosphorus availability.
increasing root area provides a larger area for the nutrient Plant and Soil 237: 225–237.
Marschner H (1995) Mineral Nutrition of Higher Plants, 2nd edn.
acquisition. Some nutrients are acquired by beneficial
New York, NY: Academic Press.
symbioses with bacteria and fungi, these relationships are
Martinoia E, Meyer S, De Angeli A and Nagy R (2012) Vacuolar
especially important in nutrient poor environments. These
transporters in their physiological context. Annual Reviews of
symbioses are important for water and nutrient acquisition Plant Biology 63: 183–213.
and there is a scope for encouraging their development Martı́nez-Granero F, Rivilla R and Martı́n M (2006) Rhizosphere
more in agricultural soils. selection of highly motile phenotypic variants of Pseudomonas
fluorescens with enhanced competitive colonization ability.
Applied and Environmental Microbiology 72: 3429–3434.
Acknowledgements Miller AJ and Smith SJ (2008) Cytosolic nitrate ion homeostasis:
could it have a role in sensing nitrogen status? Annals of Botany
Anthony J Miller is supported by grant funding (BB/ 101: 485–489.
JJ004553 and BB/JJ004561) from the BBSRC and the John Mommer L, van Ruijven J, Jansen C, van de Steeg HM and
Innes Foundation. deKroon H (2012) Interactive effects of nutrient heterogeneity
and competition: implications for root foraging theory? Func-
tional Ecology 26: 66–73.
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