Marschner’s Mineral Nutrition of Higher Plants.

© 2012 Elsevier Ltd. All rights reserved. 2012
Introduction, Definition and Classification
of Nutrients
Ernest Kirkby
Faculty of Biological Sciences, University of Leeds, UK
This chapter provides a brief overview of the history of plant
nutrition and defines the term ‘essential mineral element’, and
groups these elements according to their biochemical behaviour
and physiological functions.
The beneficial effect of adding mineral elements (e.g.,
plant ash or lime) to soils to improve plant growth has
been known in agriculture for more than 2,000 years.
Nevertheless, even 150 years ago it was still a matter of sci-
entific controversy as to whether mineral elements function
as nutrients for plant growth. It was mainly to the credit
of Justus von Liebig (1803–1873) that the scattered infor-
mation concerning the importance of certain elements for
plant growth was compiled and summarized and that the
mineral nutrition of plants was established as a scientific
discipline. These achievements led to a rapid increase in the
use of mineral fertilizers. By the end of the nineteenth cen-
tury, especially in Europe, large amounts of potash, super-
phosphate and, later, inorganic N were used in agriculture
and horticulture to improve crop growth and production.
Liebig’s conclusion that the elements N, S, P, K,
Ca, Mg, Si, Na and Fe are essential for plant growth was
reached by observation and speculation rather than by pre-
cise experimentation. The fact that the ‘mineral element
theory’ was based on this unsound foundation was one of
the reasons for the large number of studies undertaken at the
end of the nineteenth century. From these and other exten-
sive investigations on the elemental composition of differ-
ent plant species growing on various soils, it was realized
as early as the beginning of the last century that neither the
presence nor the concentration of an element in a plant is a
criterion for essentiality. Plants have a limited capability for
the selective uptake of those elements which are essential
for their growth. Additionally they take up elements which
are not needed for growth and which may even be toxic.
The elemental composition of plants growing in soils can-
not therefore be used to establish whether an element is
essential. Once this fact was appreciated, both water and
sand culture experiments were carried out in which plants
were deprived of particular elements and the consequent
effects on growth and development studied. Such investi-
gations made possible a more precise characterization of
the essentiality of elements and helped to define their role
in plant metabolism. This work also revealed two fairly
distinct groups of nutrients, the macronutrients which are
required and are present in relatively high concentrations in
plants, and the micronutrients which are equally essential,
but present in very much lower concentrations. For higher
plants, the essentiality of 14 elements is now well estab-
lished, although the requirement for the micronutrients
Cl and Ni is as yet restricted to a limited number of plant
species. Progress in this research was closely related to the
development of analytical chemistry, particularly in the
purification of chemicals and analysis. This relationship is
reflected in the time course of the discovery of the essenti-
ality of the micronutrients (Table 1.1).
The term essential mineral element (or mineral nutri-
ent) was proposed by Arnon and Stout (1939). These
authors concluded that, for an element to be considered
essential, three criteria must be met:
1. A given plant must be unable to complete its lifecycle
in the absence of the element.
Chapter 1
PART | I Nutritional Physiology 4
2. The function of the element must not be replaceable by
another element.
3. The element must be directly involved in plant metab-
olism – for example, as a component of an essen-
tial plant constituent such as an enzyme – or it must
be required for a distinct metabolic step such as an
enzyme reaction.
According to this strict definition, an element which
alleviates the toxic effects of another element (e.g., Si for
Mn toxicity), or one which simply replaces another ele-
ment (e.g., Na for K) may not be described as essential for
plant growth.
In addition to their relative concentrations within the plant,
elements may also be classified according to biochemical
behaviour and physiological function. In a scheme proposed
by Mengel and Kirkby (2001) (Table 1.2), all plant nutrients
including C, H and O as well as some non-essential ele-
ments (Si and Na) are considered. Four groups can be distin-
guished (Table 1.2).
The first group includes the major constituents of
organic plant material: C, H, O, N and S. These elements
are constituents of amino acids, proteins, enzymes and
nucleic acids, the building blocks of life. The assimilation
of all these nutrients by plants is closely linked with oxida-
tion-reduction reactions.
Phosphorus, B and Si constitute a second group of ele-
ments with close similarities in biochemical behaviour.
All three are taken up from the soil solution as inorganic
anions or acids and occur in this form in plant cells or are
bound by hydroxyl groups of sugars to form phosphate,
borate and silicate esters.
The third group of plant nutrients is made up of K,
Na, Ca, Mg, Mn and Cl, all of which are taken up from
the soil solution in the form of their ions. In plant cells,
they are also present in ionic form where they have non-
specific functions, e.g. in establishing electro-potentials.
TABLE 1.1 Discovery of the essentiality of
micronutrients for higher plants
symbol) Year Discovered by
Fe 1860 J. Sachs
Mn 1922 J.S. McHargue
B 1923 K. Warington
Zn 1926 A.L. Sommer and C.B. Lipman
Cu 1931 C.B. Lipman and G.
Mo 1938 D.I. Arnon and P.R. Stout
Cl 1954 T.C. Broyer et al.
Ni 1987 P.H. Brown et al.
TABLE 1.2 Classification of plant nutrients
Nutrient Uptake
Group 1
C, H, O, N, S as CO

O, O
, NO


, N
, SO
ions from the
soil solution, gases
from the atmosphere
Major constituents of
organic material.
Essential elements
of atomic groups
involved in enzymatic
Assimilation by
Group 2
P, B, Si as phosphates, boric
acid or borate, silic
acid from the soil
Esterification with
alcohol groups.
Phosphate esters
involved in energy
transfer reactions.
Group 3
K, Na, Ca,
Mg, Mn, Cl
as ions from the soil
functions establishing
osmotic potential.
More specific
functions for optimal
confirmation of
enzymes (enzyme
Bridging of reaction
Balancing anions.
permeability and
Group 4
Fe, Cu,
Zn, Mo
as ions or chelates
from the soil
In chelated form in
prosthetic groups of
Enable electron
transport by valency
From Mengel and Kirkby (2001) with kind permission from Springer
Science Business Media.
Chapter | 1 Introduction, Definition and Classification of Nutrients 5
The cations are associated with diffusible or indiffusible
anions, e.g. Ca with oxalate or with the carboxylic groups
of pectins in cell walls. Magnesium can also be bound
very strongly by coordinate and covalent bonds (chela-
tion) as occurs in the chlorophyll molecule. The ability of
Mg, Ca and Mn to form chelates means that these elements
closely resemble those of the fourth group, Fe, Cu, Zn and
Mo, which are predominantly present in plants in chelated
form. An important function of these latter elements is to
facilitate electron transport by valency change.
Because of continuous developments and refinements
in analytical techniques, especially in the purification of
chemicals, the current list of essential elements might well
be extended to include elements that are essential only in
very low concentrations in plants (i.e., that act as micro-
nutrients). This may possibly be the case for Na and Si,
two elements abundant in the biosphere for which essen-
tiality has already been established for some plant spe-
cies (Chapter 8). Most micronutrients are predominantly
constituents of enzyme molecules and are thus essential
only in small amounts at the whole plant level. By con-
trast, the macronutrients are either constituents of organic
compounds, such as proteins and nucleic acids, or act as
osmotica. These differences in function are reflected in the
average concentrations of mineral nutrients in plant shoots
that are sufficient for adequate growth (Table 1.3). The
values can vary considerably depending on plant species,
plant age, and concentration of other mineral elements.
This aspect is discussed in Chapters 6 to 8.
TABLE 1.3 Average concentrations of mineral elements
in plant shoot dry matter sufficient for adequate
symbol µmol g
dw mg kg
Molybdenum Mo 0.001 0.1
Nickel Ni 0.001 0.1
Copper Cu 0.1 6
Zinc Zn 0.3 20
Manganese Mn 1.0 50
Iron Fe 2.0 100
Boron B 2.0 20
Chlorine Cl 3.0 100
Sulphur S 30 1,000
Phosphorus P 60 2,000
Magnesium Mg 80 2,000
Calcium Ca 125 5,000
Potassium K 250 10,000
Nitrogen N 1,000 15,000
From Epstein (1965), Epstein and Bloom (2005), Brown et al. (1987b).

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