Plant Mineral Nutrition

Plants & Inorganic Nutrients: Objectives

This chapter will examine the nutritional requirements of

plants that are satisfied by mineral elements.

• methods employed in the study of mineral nutrition.

• concept of essential and beneficial elements and

distinction between macronutrients and
micronutrients.

• discussion of the metabolic roles of the essential

mineral elements, including the concept of critical and
deficient concentration, and symptoms that are
associated with deficiencies of the mineral elements.

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Roots, Soils, & Nutrient Uptake: Objectives

This chapter will also examine the availability of nutrients in the soil
and their uptake by roots.

• soil as a source of nutrient elements, the colloidal nature of soil,

and ion exchange properties that determine the availability of
nutrient elements in a form that can be taken up by roots.

• mechanism of solute transport across membranes: simple and

facilitated diffusion; active transport; functions of membrane
proteins as ion channels and carriers; role of electrochemical
gradients.

• ion traffic into and through the root tissues and the concept of
apparent free space.

• beneficial role of microorganisms, especially fungi, with respect

to nutrient uptake by roots.

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 CO2 H2O

O2

Plants derive most of their

organic mass from the CO2
of air, but they also depend
on soil nutrients, such as
water and minerals.

O2
Minerals

CO2

H2O
 Plants are immobile and autotrophic.

• Plants must obtain the nutrients they need to

live from the medium in which they grow - soil is
a plant's supermarket.

• Plants are the primary agents of mineral

incorporation into the biosphere - they are
biology's miners.
What does a plant need to survive and grow?

How do we figure out which ones are essential

for plant growth and reproduction?

Try to grow plants in a precisely defined

medium, then remove individual components
and observe the effect on growth.

Do this using hydroponics - growing plants in

a liquid medium, with precise concentrations of
elements added to absolutely pure water.
Control: Solution Experimental: Solution
containing all minerals without potassium
 Special Techniques Used in Nutritional Studies

• The technique of growing plants with their roots

immersed in nutrient solution without soil is called
solution culture or hydroponics.
 Essential Elements: Criteria

1) Required for normal growth and completion

of the life cycle

2) Not replaceable under normal growth

conditions

3) Have a demonstrated biological role in the plant

- structural or physiological function
Macronutrients and Micronutrients
 Group 1 - Nutrients that are part of carbon
compounds

N Constituent of amino acids, amides, proteins, nucleic

acids, nucleotides, coenzymes, hexoamines, etc.

S Component of cysteine, cystine, methionine, and

proteins.
Constituent of lipoic acid, coenzyme A, thiamine
pyrophosphate, glutathione, biotin, adenosine-5′-
phosphosulfate, and 3-phosphoadenosine.
Group 2 - Nutrients that are important in energy
storage or structural integrity
P Component of sugar phosphates, nucleic acids,
nucleotides, coenzymes, phospholipids, phytic acid, etc.
Has a key role in reactions that involve ATP.

Si Deposited as amorphous silica in cell walls.

Contributes to cell wall mechanical properties,
including rigidity and elasticity.

B Complexes with mannitol, mannan,

polymannuronic acid, and other constituents of cell walls.
Involved in cell elongation and nucleic acid
metabolism.
 Group 3 Nutrients that remain in ionic form
(enzyme cofactors and regulators of osmotic potentials)

K Required as a cofactor for more than 40 enzymes.

Principal cation in establishing cell turgor and
maintaining cell electroneutrality.

Ca Constituent of the middle lamella of cell walls.

Required as a cofactor by some enzymes involved in
the hydrolysis of ATP and phospholipids.
Acts as a second messenger in metabolic regulation.

Mg Required by many enzymes involved in phosphate

transfer.
Constituent of the chlorophyll molecule.
 Group 3 (cont.)

Cl Required for the photosynthetic reactions involved in

O2 evolution.

Mn Required for activity of some dehydrogenases,

decarboxylases, kinases, oxidases, and peroxidases.
Involved with other cation-activated enzymes and
photosynthetic O2 evolution.

Na Involved with the regeneration of phosphoenol-

pyruvate in C4 and CAM plants.
Substitutes for potassium in some functions.
 Group 4 - Nutrients that are involved in redox
reactions
Fe Constituent of cytochromes and nonheme iron
proteins involved in photosynthesis, N2 fixation, and
respiration.

Zn Constituent of alcohol dehydrogenase, glutamic

dehydrogenase, carbonic anhydrase, etc.

Cu Component of ascorbic acid oxidase, tyrosinase,

monoamine oxidase, uricase, cytochrome oxidase,
phenolase, laccase, and plastocyanin.

Ni Constituent of urease.
In N2-fixing bacteria, constituent of hydrogenases.

Mo Constituent of nitrogenase, nitrate reductase, and

xanthine dehydrogenase.
 Beneficial elements

Na, Co - required by particular plants

• Na and Co are taken up by plants - are

generally not required - BUT are required
in the nutrition of the animals or micro-
organisms that feed on them.

• Co – essential for nitrogen-fixing bacteria,

associated with legumes.
 Mobile and Immobile Elements
• some elements can be remobilized by the plant
=> mobile elements

• some are fixed in location once they are

deposited => immobile elements

Immobile elements - Ca, S, Fe, Mn, B, Cu

• Ca is locked into the cell walls
• metals are not easily metabolized or transported.
• deficiency symptoms seen in young leaves.

Mobile elements - N, P, K, Mg, Cl, Na, Zn, Mo

• N, P can be metabolized easily; K, Mg quite mobile
• deficiency seen first in older leaves.
 Symptoms of Mineral Deficiency
What happens when there is a deficiency in a nutrient?
• Symptoms of mineral deficiency depend on the nutrient’s function and
mobility within the plant.
• Since we know at the molecular level how the elements function, we can begin
to predict some of the effects of a deficiency in that element.
• Vice versa, deficiency symptoms of an element begin to give clues about its
function.
• Deficiency of a mobile nutrient usually affects older organs more than young
ones.
• Deficiency of a less mobile nutrient usually affects younger organs more
than older ones.
• The most common deficiencies are those of nitrogen, potassium, and
phosphorus –kaya fertilizers have these stuff.
Healthy

Phosphate-deficient

Potassium-deficient

Nitrogen-deficient
Mineral Deficiencies Disrupt Plant Metabolism
and Function

General Deficiency Symptoms

• Stunted growth

• Chlorosis

• Necrosis
 Relationship between yield (or growth) and the
nutrient content of the plant tissue.

• The effect of varying the concentration of a nutrient during plant growth is

reflected in the growth or yield.
• The yield parameter may be expressed in terms of shoot dry weight or
height.
• Three zones of nutrient concentration —deficiency, adequate, and toxic.
• The critical concentration for that nutrient is the concentration at which
below it, yield or growth is reduced.
Mineral Uptake
 Mineral Uptake: Key Points

• Mineral movement to root by diffusion or

bulk flow and root growth.
• Uptake controlled at root endodermis.
• Uptake by either:
-- simple diffusion (no protein),
-- facilitated diffusion (carrier or channel), or
-- active uptake (requires energy and a
protein carrier).
 Negatively Charged Soil Particles Affect the Absorption
of Mineral Nutrients into the Root Tissues.

Soil particle surrounded by

film of water
Soil particle
Root hair

Water
available
to plant

Root hair

Air space

Soil water
Cation exchange in soil

Cation exchange capacity (CEC) - the degree to which a soil can

adsorb and exchange ions.
• Acids derived from roots contribute to a plant’s
uptake of minerals when H+ displaces mineral
cations from clay particles.
 Influence of soil pH on the availability of nutrient
elements in organic soils.

• The width of the shaded areas indicates the degree of nutrient availability
to the plant root.
• All of these nutrients are available in the pH range of 5.5 to 6.5.
 Soil pH Affects Growth of Soil Microbes
and Roots, & Nutrient Solubility

• Hydrogen ion concentration (pH) is an important property of soils

because it affects the growth of plant roots and soil
microorganisms.

• Root growth is generally favored in slightly acidic soils, at pH values

between 5.5 and 6.5.

• Fungi generally predominate in acidic soils; bacteria become more

prevalent in alkaline soils.

• Soil pH affects the solubility of soil nutrients.

Acidity promotes the weathering of rocks that releases K+, Mg2+,

Ca2+, and Mn2+ and increases solubility of carbonates, sulfates, and
phosphates. Increasing the solubility of nutrients facilitates their
availability to roots.
 Plants Develop Extensive Root Systems

Fibrous root systems of wheat (a monocot).

(A) The root system of a mature (3-month-old) wheat plant
growing in dry soil.
(B) The root system of a wheat plant growing in irrigated soil.
Taproot system of two adequately watered dicots: sugar beet and alfalfa.
• The root system shows a major vertical root axis. In the case of sugar
beet, the upper portion of the taproot system is thickened because of
its function as storage tissue.
 Different Areas of the Root Absorb Different
Mineral Ions
The high rates of nutrient absorption in the apical root zones due to:
-- the strong demand for nutrients in these tissues
- the relatively high nutrient availability in the soil surrounding them.

• Root absorption of calcium in barley appears to be restricted to the apical

region.
• Iron may be taken up either at the apical region, as in barley (Clarkson 1985),
or over the entire root surface, as in corn (Kashirad et al. 1973).
• Potassium, nitrate, ammonium, and phosphate can be absorbed freely at
all locations of the root surface (Clarkson 1985), but in corn the elongation
zone has the maximum rates of potassium accumulation (Sharp et al. 1990)
and nitrate absorption (Taylor and Bloom 1998).
• In corn and rice, the root apex absorbs ammonium more rapidly than the
elongation zone does (Colmer and Bloom 1998).
• In several species, root hairs are the most active in phosphate absorption
(Fohse et al. 1991).
 Concept of Apparent Free Space
• AFS – a fraction of the root tissue volume that is
not separated from the environment by a
membrane or other diffusion barrier.
• What constitutes the AFS of the root?
cell walls and intercellular spaces (= apoplast of
the epidermis and cortex)
• It comprises the volume of the root that can be
entered without crossing a barrier, or that is
accessible by free diffusion.
• CEC of the AFS is due to the carboxyl groups
associated with galacturonic acid residues in the
CW pectic compounds.
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Plants & Nitrogen
 Nitrogen is often the mineral that has the greatest
effect on plant growth.

• Plants require nitrogen as a component of

proteins, nucleic acids, chlorophyll, and other
important organic molecules
 Soil Bacteria and Nitrogen Availability
• Nitrogen fixation - converts atmospheric N2 to nitrogenous
compounds (NH3 and NO- 3) that plants can absorb as a
nitrogen source for organic synthesis.

• Nitrogen Cycle
Ammonification - bacterial and fungal catabolism
soil organic matter  NH4+.
Nitrification - Nitrosomonas sp.: NH4+ NO2-
and

- Nitrobacter sp.: NO2-  NO3-

Denitrification - bacterial conversion

NO3-  NO  N2
Nitrogen Cycle
 Plant nutritional adaptations often involve
relationships with other organisms

• Two types of relationships plants have with other

organisms are mutualistic:

– Symbiotic nitrogen fixation, involving roots and

bacteria

– Mycorrhizae, involving roots and fungi

The Role of Bacteria in Symbiotic Nitrogen Fixation

• Symbiotic relationships with nitrogen-fixing

bacteria provide some plant species with a built-in
source of fixed nitrogen.

• The key symbioses between plants and nitrogen-

fixing bacteria occur in the legume family
(peas, beans, and other similar plants)
 5 µm

Bacteroids
within
vesicle

Nodules

Roots

Pea plant root. Bacteroids in a soybean

root nodule.
• Along a legume’s roots are swellings called
nodules, composed of plant cells ―infected‖ by
nitrogen-fixing Rhizobium bacteria.
• Inside the root nodule, Rhizobium bacteria
assume a form called bacteroids, which are
contained within vesicles formed by the root cell.
• The bacteria of a root nodule obtain sugar from
the plant and supply the plant with fixed
nitrogen.

• Each legume species is associated with a

particular strain of Rhizobium.
 Rhizobium
bacteria
Infection Dividing cells
thread in root cortex
Bacteroid Dividing cells in
Infected pericycle
root hair

Developing
root nodule

Bacteroid

Nodule
vascular
Bacteroid tissue
 Mycorrhizae and Plant Nutrition

Mycorrhizal Fungi Facilitate Nutrient Uptake by Roots

• Mycorrhizae are mutualistic associations of fungi

and roots.

• The fungus benefits from a steady supply of sugar

from the host plant.

• The host plant benefits because the fungus

increases the surface area for water uptake and
mineral absorption.
Formation of a nutrient depletion zone in the
region of the soil adjacent to the plant root.

Nutrient depletion zone -- forms when the rate of nutrient uptake by

the cells of the root exceeds the rate of replacement of the nutrient
by diffusion in the soil solution. This depletion causes a localized
decrease in the nutrient concentration in the area adjacent to the
root surface.
Two Main Types of Mycorrhizae

• ectomycorrhizae

• endomycorrhizae
In ectomycorrhizae, the mycelium of the fungus forms
a dense sheath over the surface of the root.
Epidermis Cortex Mantle 100 µm
(fungal
sheath)

Endodermis

Fungal
hyphae
between
Mantle cortical
(fungal sheath) cells
(colorized SEM)
Ectomycorrhizae.
In endomycorrhizae, microscopic fungal hyphae
extend into the root.
Epidermis Cortex 10 µm
Cortical cells

Endodermis

Fungal
Vesicle
hyphae
Casparian
strip
Root
hair Arbuscules

(LM, stained specimen)

Endomycorrhizae.
 Root infected with ectotrophic mycorrhizal fungi

In the infected root, the fungal hyphae surround the root to produce a dense fungal
sheath and penetrate the intercellular spaces of the cortex to form the
Hartig net.
The total mass of fungal hyphae may be comparable to the root mass itself.
Association of vesicular-arbuscular mycorrhizal
fungi with a section of a plant root.

• The fungal hyphae grow into the intercellular wall spaces of the cortex and
penetrate individual cortical cells.
• As they extend into the cell, they do not break the plasma membrane or the
tonoplast of the host cell. Instead, the hypha is surrounded by these
membranes and forms structures known as arbuscules and vesicles, which
participate in nutrient ion exchange between the host plant and the fungus.
 Quiz
Discuss the importance of the following in nutrient
uptake in plants:
1. soil pH
2. cation-exchange capacity (CEC)
3. root growth
4. microorganisms

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings