Part I

Nutritional Physiology
Copyright 2011. Elsevier Science. All rights reserved.

Marschner, Horst. Marschner's Mineral Nutrition of Higher Plants, Elsevier Science, 2011. ProQuest Ebook Central, http://ebookcentral.proquest.com/lib/sibdilibro-ebooks/detail.action?docID=858643.
Created from sibdilibro-ebooks on 2017-09-05 11:31:20.
 This page intentionally left blank
Copyright 2011. Elsevier Science. All rights reserved.

Marschner, Horst. Marschner's Mineral Nutrition of Higher Plants, Elsevier Science, 2011. ProQuest Ebook Central, http://ebookcentral.proquest.com/lib/sibdilibro-ebooks/detail.action?docID=858643.
Created from sibdilibro-ebooks on 2017-09-05 11:31:20.

Introduction, Definition and Classification
of Nutrients
Ernest Kirkby
Faculty of Biological Sciences, University of Leeds, UK
SUMMARY
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.
1.1 GENERAL
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 (18031873) 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-
Copyright 2011. Elsevier Science. All rights reserved.
phosphate and, later, inorganic N were used in agriculture
and horticulture to improve crop growth and production.
Liebigs 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
Marschners Mineral Nutrition of Higher Plants. DOI: 10.1016/B978-0-12-384905-2.00001-7
Marschner, Horst. Marschner's Mineral
2012 Elsevier Nutrition
Ltd. All of Higher Plants, Elsevier Science, 2011. ProQuest Ebook Central, http://ebookcentral.proquest.com/lib/sibdilibro-ebooks/detail.action?docID=858643.
rights reserved.
Created from sibdilibro-ebooks on 2017-09-05 11:31:20.
Chapter 1
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.
1.2 ESSENTIAL ELEMENTS FOR PLANT
GROWTH
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.
3
 4 PART | I Nutritional Physiology

TABLE 1.1 Discovery of the essentiality of TABLE 1.2 Classification of plant nutrients
micronutrients for higher plants
Biochemical
Element Nutrient Uptake functions
(chemical Group 1
symbol) Year Discovered by
C, H, O, N, S as CO2, HCO3, Major constituents of
Fe 1860 J. Sachs organic material.
H2O, O2, NO3,
Mn 1922 J.S. McHargue NH4, N2, SO42, Essential elements
SO2 ions from the of atomic groups
B 1923 K. Warington soil solution, gases involved in enzymatic
Zn 1926 A.L. Sommer and C.B. Lipman from the atmosphere processes.
Assimilation by
Cu 1931 C.B. Lipman and G. oxidation-reduction
MacKinney reactions.
Mo 1938 D.I. Arnon and P.R. Stout Group 2
Cl 1954 T.C. Broyer et al. P, B, Si as phosphates, boric Esterification with
acid or borate, silic alcohol groups.
Ni 1987 P.H. Brown et al.
acid from the soil Phosphate esters
solution involved in energy
transfer reactions.
Group 3
2. The function of the element must not be replaceable by
another element. K, Na, Ca, as ions from the soil Non-specific
Mg, Mn, Cl solution functions establishing
3. The element must be directly involved in plant metab-
osmotic potential.
olism for example, as a component of an essen- More specific
tial plant constituent such as an enzyme or it must functions for optimal
be required for a distinct metabolic step such as an confirmation of
enzyme reaction. enzymes (enzyme
activation).
According to this strict definition, an element which Bridging of reaction
alleviates the toxic effects of another element (e.g., Si for partners.
Balancing anions.
Mn toxicity), or one which simply replaces another ele-
Controlling
ment (e.g., Na for K) may not be described as essential for membrane
plant growth. permeability and
electrochemical
potentials.
1.3 BIOCHEMICAL BEHAVIOUR AND
Group 4
PHYSIOLOGICAL FUNCTIONS OF
ELEMENTS IN PLANTS Fe, Cu, as ions or chelates In chelated form in
Zn, Mo from the soil prosthetic groups of
In addition to their relative concentrations within the plant, solution enzymes.
elements may also be classified according to biochemical Enable electron
Copyright 2011. Elsevier Science. All rights reserved.

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
transport by valency
change.
From Mengel and Kirkby (2001) with kind permission from Springer
Science Business Media.
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.

Marschner, Horst. Marschner's Mineral Nutrition of Higher Plants, Elsevier Science, 2011. ProQuest Ebook Central, http://ebookcentral.proquest.com/lib/sibdilibro-ebooks/detail.action?docID=858643.
Created from sibdilibro-ebooks on 2017-09-05 11:31:20.
 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 TABLE 1.3 Average concentrations of mineral elements
of pectins in cell walls. Magnesium can also be bound in plant shoot dry matter sufficient for adequate
very strongly by coordinate and covalent bonds (chela- growth
tion) as occurs in the chlorophyll molecule. The ability of Chemical
Mg, Ca and Mn to form chelates means that these elements Element symbol mol g1 dw mg kg1
closely resemble those of the fourth group, Fe, Cu, Zn and
Molybdenum Mo 0.001 0.1
Mo, which are predominantly present in plants in chelated
form. An important function of these latter elements is to Nickel Ni 0.001 0.1
facilitate electron transport by valency change. Copper Cu 0.1 6
Because of continuous developments and refinements Zinc Zn 0.3 20
in analytical techniques, especially in the purification of
chemicals, the current list of essential elements might well Manganese Mn 1.0 50
be extended to include elements that are essential only in Iron Fe 2.0 100
very low concentrations in plants (i.e., that act as micro- Boron B 2.0 20
nutrients). This may possibly be the case for Na and Si,
Chlorine Cl 3.0 100
two elements abundant in the biosphere for which essen-
tiality has already been established for some plant spe- Sulphur S 30 1,000
cies (Chapter 8). Most micronutrients are predominantly Phosphorus P 60 2,000
constituents of enzyme molecules and are thus essential
Magnesium Mg 80 2,000
only in small amounts at the whole plant level. By con-
trast, the macronutrients are either constituents of organic Calcium Ca 125 5,000
compounds, such as proteins and nucleic acids, or act as Potassium K 250 10,000
osmotica. These differences in function are reflected in the
Nitrogen N 1,000 15,000
average concentrations of mineral nutrients in plant shoots
that are sufficient for adequate growth (Table 1.3). The From Epstein (1965), Epstein and Bloom (2005), Brown et al. (1987b).
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.
Copyright 2011. Elsevier Science. All rights reserved.

Marschner, Horst. Marschner's Mineral Nutrition of Higher Plants, Elsevier Science, 2011. ProQuest Ebook Central, http://ebookcentral.proquest.com/lib/sibdilibro-ebooks/detail.action?docID=858643.
Created from sibdilibro-ebooks on 2017-09-05 11:31:20.
 This page intentionally left blank
Copyright 2011. Elsevier Science. All rights reserved.

Marschner, Horst. Marschner's Mineral Nutrition of Higher Plants, Elsevier Science, 2011. ProQuest Ebook Central, http://ebookcentral.proquest.com/lib/sibdilibro-ebooks/detail.action?docID=858643.
Created from sibdilibro-ebooks on 2017-09-05 11:31:20.