ve in an entirely inorganic environment.

As
autotrophic organisms, plants must take in carbon
dioxide from the atmosphere and water and mineral
nutrients from the soil, and from these simple, inorganic
components, make all of the complex molecules of
a living organism. Since plants stand at the bottom
of the food chain, mineral nutrients assimilated by
plants eventually find their way into the matter that
makes up all animals, including humans. All organisms
must continually draw material substance from their
environment in order to maintain their metabolism,
growth, and development. The means for making these
materials available to the organism is the subject of
plant nutrition.
Plant nutrition is traditionally treated as two separate topics: organic nutrition and inorganic nutrition.
Organic nutrition focuses on the production of carbon
compounds, specifically the incorporation of carbon,
hydrogen, and oxygen via photosynthesis, while inorganic nutrition is concerned primarily with the acquisition
of mineral elements from the soil. Photosynthesis
and the acquisition of mineral ions from the soil are so
interdependent, however, that this distinction between
organic and inorganic nutrition is more a matter of convenience than real. Nevertheless, because the acquisition
and assimilation of carbon are addressed in Chapters 7
and 8, this chapter will focus on the acquisition of mineral elements and the role of those elements in plant
metabolism.
This chapter will examine the nutritional requirements of plants that are satisfied by mineral elements.
This will include
• methods employed in the study of mineral nutrition,
• the concept of essential and beneficial elements
and the distinction between macronutrients and
micronutrients,
• a general discussion of the metabolic roles of the 14
essential mineral elements, the concept of critical
and deficient concentration, and symptoms associated with deficiencies of the mineral elements,
and
• a brief discussion of micronutrient toxicity.
The soil as a nutrient reservoir and mechanisms for
mineral uptake by roots was covered in Chapter 3.
61
62 Chapter 4 / Plants and Inorganic Nutrients
4.1 METHODS AND NUTRIENT
SOLUTIONS
4.1.1 INTEREST IN PLANT NUTRITION
IS ROOTED IN THE STUDY OF
AGRICULTURE AND CROP
PRODUCTIVITY
Much of the groundwork for modern nutritional studies
was laid in Europe in the early to mid-nineteenth century, in response to a combination of political and social
factors. The Napoleonic wars had devastated Europe
and the industrial revolution was gaining momentum.
Rising populations and massive migration to the cities
created demands that could no longer be met by the
traditional agricultural economy, one that relied heavily on the use of organic manures. Greater efficiency
in agriculture was required and this was not possible without a more thorough understanding of plant
nutrition.
One of the first to make significant progress in
the study of plant nutrition was N. T. de Saussure
(1767–1845), who studied both photosynthesis and the
absorption of nutrient elements with the same careful, quantitative methods. De Saussure conducted some
of the first elemental analyses of plant material and
introduced the concept that some, but not necessarily
all, of the elements found might be indispensable, or
essential, to plant growth. De Saussure’s ideas concerning the importance of elements derived from the soil
generated considerable debate at the time, but received
support from the work of C. S. Sprengel (1787–1859),
working in Germany, and Jean-Baptiste Boussingault in
France. Sprengel introduced the idea that soils might
be unproductive if deficient in but one single element
necessary for plant growth, and Boussingault stressed
quantitative relationships between the effects of fertilizer and nutrient uptake on crop yields. Boussingault
is also credited with providing the first evidence that
legumes had the unique capacity to assimilate atmospheric nitrogen, a finding that was later confirmed by
the discovery of the nitrogen-fixing role of bacteria in
root nodules.
By the middle of the nineteenth century, many
pieces of the nutritional puzzle were beginning to fall
into place. In 1860, Julius Sachs, a prominent German
botanist, demonstrated for the first time that plants could
be grown to maturity in defined nutrient solutions in the
complete absence of soil. J. B. Lawes and J. H. Gilbert,
working at Rothamsted in England, had successfully
converted insoluble rock phosphate to soluble phosphate (called superphosphate), and by the end of the
century the agricultural use of NPK (nitrogen, phosphorous, and potassium) fertilizers was well established in
Europe.
4.1.2 THE USE OF HYDROPONIC
CULTURE HELPED TO DEFINE
THE MINERAL REQUIREMENTS
OF PLANTS
In the mid-nineteenth century, J. Sachs was interested
in determining the minimal nutrient requirements of
plants. Recognizing that it would be difficult to pursue
such studies in a medium as complex as soil, Sachs
devised an experimental system such that the roots
grew not in soil but in an aqueous solution of mineral
salts. With this simplified system, Sachs was able to
demonstrate the growth of plants to maturity on a relatively simple nutrient solution containing six inorganic
salts (Table 4.1). Variations on Sachs’s system, known
as solution or hydroponic culture (growing plants in
a defined nutrient solution), have remained to this day
the principal experimental system for study of plant
nutrient requirements. Hydroponic culture is also now
used extensively in North America for the year-round
commercial production of vegetables such as lettuce,
tomato, sweet peppers, and seedless cucumber.
The nutrient solution devised by Sachs contributed
a total of nine mineral nutrients (K, N, P, Ca, S, Na, Cl,
Fe, Mg). Carbon, hydrogen, and oxygen were excluded
from this total because they were provided in the form
of carbon dioxide and water and were not considered
mineral elements. It was at least another half century
before the need for additional mineral nutrients was
demonstrated. There was no magic to the success of
Sachs’s experiments. Many of the mineral nutrients used
by plants are required in very low amounts and Sachs
unknowingly provided these nutrients as impurities in
the salts and water he used to make up his nutrient solution. Analytical techniques have now improved
to the point where it is possible to detect mineral
contents several orders of magnitude lower than was
possible in Sachs’s time. Most mineral elements are
now measured by either atomic absorption spectrometry or atomic emission spectrometry. These techniques
involve vaporization of the elements at temperatures
TABLE 4.1 The composition of Sachs’s nutrient
solution (1860) used for solution culture of plants.
Approximate
Concentration
Salt Formula (mM)

Potassium nitrate KNO3 9.9

Calcium phosphate Ca3(PO4)2 1.6
Magnesium sulfate MgSO4z7H2O 2.0
Calcium sulfate CaSO4 3.7
Sodium chloride NaCl 4.3

Iron sulfate FeSO4 trace

4.1 Methods and Nutrient Solutions 63

in excess of several thousand degrees. In the vaporous
state, the element will either absorb or emit light at
very narrow wavelength bands. The wavelength of light
absorbed or emitted is characteristic of a particular element and the quantity of absorbed or emitted energy
is proportional to the concentration of the element
in the sample. In this way, concentrations as low as
10-8 g ml-1 for some elements can be measured in
samples of plant tissue, soil, or nutrient solutions within
a few minutes.
Aside from the commercial applications of
hydroponic plant culture, a great deal of plant
physiology and other botanical research is conducted
with plants grown under controlled environments. This
may include relatively simple greenhouses or complex
growth rooms in which temperature and lighting are
carefully regulated. Plant nutrient supply must also be
regulated, and over the years a large number of nutrient
solutions have been formulated for this purpose. Most
modern formulations are based on a solution originally
developed by D. R. Hoagland, a pioneer in the study
of plant mineral nutrition. Individual investigators
may introduce minor modifications to the composition of the nutrient solution in order to accommodate specific
needs. Such formulations are commonly
referred to as modified Hoagland’s solutions
(Tables 4.2, 4.3).
The concentration of minerals in most nutrient
solutions is many times greater than that normally found
in soils. An excess is necessary in order to maintain a
continual supply of nutrients as they are taken up by the
roots. The nutrient concentration of the soil solution,
on the other hand, is relatively low but is continually
replenished by nutrients adsorbed on the soil particles
(Chapter 3).
TABLE 4.2 The composition of a typical
one-half strength ‘‘modified’’ Hoagland’s nutrient
solution, showing the nutrient salts used and their
approximate millimolar (mM) concentrations.
Concentration
(mM)
Calcium nitrate Ca(NO)3 2.5
Potassium phosphate KH2PO4 0.5
Potassium nitrate KNO3 2.5
Magnesium sulfate MgSO4 1.0
Zinc sulfate ZnSO4 0.00039
Manganous sulfate MnSO4 0.0046
Copper sulfate CuSO4 0.00016
Boric acid H3BO3 0.0234
Molybdic acid MoO3 0.000051
Iron sequestrene Fe 0.179

TABLE 4.3 The quantity of each nutrient

element in modified Hoagland’s nutrient solution.

Element Mg/L

Calcium 103

Nitrogen 105

Potassium 118

Sulfur 33

Magnesium 25

Phosphorous 15

Iron 10
Boron 0.25
Manganese 0.25
Zinc 0.025
Copper 0.01
Molybdenum 0.0052

4.1.3 MODERN TECHNIQUES

OVERCOME INHERENT
DISADVANTAGES OF SIMPLE
SOLUTION CULTURE
In the simplest form of solution culture, a seedling is
supported in the lid of a container, with its roots free
to grow in the nutrient solution (Figure 4.1). Note that
the solution must be aerated in order to obtain optimal
root growth and nutrient uptake. A solution that is not
aerated becomes depleted of oxygen, a condition known
as anoxia. Anoxia inhibits the respiration of root cells
and, because nutrient uptake requires energy, reduces
nutrient upt