1

Introduction, Definition, and Classification

of Mineral Nutrients

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 2000 years.
Nevertheless, even 150 years ago it was still a matter of scientific 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 information concerning the
importance of mineral elements for plant growth was compiled and summarized and
that the mineral nutrition of plants was estabhshed as a scientific discipline. These
achievements led to a rapid increase in the use of mineral fertilizers. By the end of the
nineteenth century, especially in Europe, large amounts of potash, superphosphate,
and, later, inorganic nitrogen were used in agriculture and horticulture to improve plant
growth.
Liebig's conclusion that the mineral elements nitrogen, sulfur, phosphorus, potass-
ium, calcium, magnesium, silicon, sodium, and iron are essential for plant growth was
arrived at by observation and speculation rather than by precise 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 extensive investigations on the mineral composition of
different plant species growing on various soils, it was reaUzed as early as the beginning
of this century that neither the presence nor the concentration of a mineral element in a
plant is a criterion for essentiality. Plants have a limited capability for the selective
uptake of those mineral elements which are essential for their growth. They also take up
mineral elements which are not necessary for growth and which may even be toxic.
The mineral composition of plants growing in soils cannot therefore be used to
estabhsh whether a mineral element is essential. Once this fact was appreciated, both
water and sand culture experiments were carried out in which particular mineral
elements were omitted. These techniques made possible a more precise characteriz-
ation of the essentiality of mineral elements and led to a better understanding of their
role in plant metabohsm. Progress in this research was closely related to the develop-
ment of analytical chemistry, particularly in the purification of chemicals and methods
of estimation. This relationship is reflected in the time scale of the discovery of the
essentiality of micronutrients (Table 1.1).
 Mineral Nutrition of Higher Plants

Table 1.1
Discovery of the Essentiality of Micronutrients for Higher Plants
Element Year Discovered by

Iron 1860 J. Sachs

Manganese 1922 J. S. McHargue
Boron 1923 K. Warington
Zinc 1926 A. L. Sommer and C. B. Lipman
Copper 1931 C. B. Lipman and G. MacKinney
Molybdenum 1938 D. I. Amon and P. R. Stout
Chlorine 1954 T. C. Broyer et al.
Nickel 1987 P. H. Brown et al.

The term essential mineral element (or mineral nutrient) 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 Hfe cycle in the absence of the mineral
element.
2. The function of the element must not be replaceable by another mineral element.
3. The element must be directly involved in plant metabolism - for example, as a
component of an essential plant constituent such as an enzyme - or it must be required
for a distinct metaboUc step such as an enzyme reaction.
According to this strict definition those mineral elements which compensate for the
toxic effects of other elements or which simply replace mineral nutrients in some of their
less specific functions, such as maintenance of osmotic pressure, are not essential, but
can be described as 'beneficial' elements (Chapter 10). It is still difficult to generalize
when discussing which mineral elements are essential for plant growth. This is
particularly obvious when higher and lower plants are compared (Table 1.2). For higher
plants the essentiaUty of 14 mineral elements is well estabhshed, although the known
requirement for chlorine and nickel is as yet restricted to a limited number of plant
species.

Table 1.2
Essentiality of Mineral Elements for Higher and Lower Plants
Classification Element Higher plants Lower plants
Macronutrient N, P, S, K, Mg, Ca + 4- (Exception: Ca for fungi)
Micronutrient Fe, Mn, Zn, Cu, + + (Exception: B for fungi)
B, Mo, CI, Ni
Micronutrient and Na, Si, Co ± ±
'beneficial' element i,v — ±

Because of continuous improvements in analytical techniques, especially in the

purification of chemicals, this list might well be extended to include mineral elements
that are essential only in very low concentrations in plants (i.e., that act as micro-
nutrients). This holds true in particular for sodium and silicon, which are abundant in
Introduction, Definition, and Classification of Mineral Nutrients 5

the biosphere. The essentiality of these two mineral elements has been established for
some higher plant species (Chapter 10). Most micronutrients are predominantly
constituents of enzyme molecules and are thus essential only in small amounts. In
contrast, the macronutrients either are 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 8 to 10,

Table 1.3
Average Concentrations of Mineral Nutrients in Plant Shoot Dry Matter that are Sufficient for
Adequate Growth"*
//molg ^ mgkg ^ Relative number
Element Abbreviation dry wt (ppm) % of atoms

Molybdenum Mo 0.001 0.1 1

Nickel^ Ni -0.001 -0.1 — 1
Copper Cu 0.10 6 — 100
Zinc Zn 0.30 20 — 300
Manganese Mn 1.0 50 — 1000
Iron Fe 2.0 100 — 2000
Boron B 2.0 20 — 2000
Chlorine CI 3.0 100 — 3000
Sulfur S 30 — 0.1 30000
Phosphorus P 60 — 0.2 60000
Magnesium Mg 80 — 0.2 80000
Calcium Ca 125 — 0.5 125000
Potassium K 250 — 1.0 250000
Nitrogen N 1000 — 1.5 1000000

^From Epstein (1965).

^Based on Brown et al. (1987b).