THEME 3: MINERAL NUTRITION

Mineral nutrition of plants covers a wide field:

- Nutrient requirements
- Functions of mineral nutrients
- Interactions between nutrients
- Root absorption of mineral nutrients
- Transport in the plant.
Mineral nutrients are elements such as N, P, K acquired primarily in the form of inorganic
ions from the soil. These elements remain as ash after combustion of dry plant material
in the laboratory.
The analysis of plant tissues reveals the presence of elements other than C, O and H
(which can be obtained from CO2 and water). Only a fraction of these mineral nutrients
are considered essential. For an element to be considered essential, three criteria must be
met:
- A given plant must be unable to complete its life cycle in the absence of the
mineral element. Therefore, the essentiality of some elements has only been
established for some species, this could be the case of Na or Si.
- The function of the element must not be replaceable by another mineral element.
- The element must be directly involved in the plant metabolism (the sum of all the
chemical reactions that take place in the cells of a living organism. These reactions
are catalyzed by enzymes. Some essential nutrients are required as a cofactor by
several enzymes. Others are constituents of many enzyme molecules).
Essential nutrients are classified according to their
relative concentration in plant tissues:
- Macronutrients: > 1 mg/g dry matter: N P K Ca
Mg S.
- Micronutrient: < 0.1 mg/g dry matter: Fe Cl B Mn
Zn Cu Mo
The current concentration of each nutrient can vary
considerably de depending on plant species, age and
organ. Nutrient concentration is higher in leaves then in
shoot (branches and leaves (these have higher concentrations due to their organs)) or
roots.
FUNCTIONS OF THE ESSENTIAL NUTRIENTS (examples)
- N, S: constituent of amino acids (proteins), S (cistine).
- P: constituent of phospholipids, nucleotides, ATP.
- Mg: constituent of the chlorophyll molecule.
- Mn: involved in photosynthetic O2 evolution.
- B, Ca: required for cell wall structural integrity.
- Ca: second messenger and membrane stability, if there is not enough the
plasmomembrane starts to behave like a cloudy barrier and not as an specific one,
this is why there are not much Ca in the cytosol.
- K, Cl: osmoregulation (changes in the turgor of the cell), cofactor > 40 enzymes.
- Fe, Zn, Cu, Mo: involved in redox reactions.
INTERACTIONS BETWEEN ESSENTIAL NUTRIENTS
Interactions between nutrients in higher plants occur when the supply
of one nutrient affects the absorption, distribution, or function of
another nutrient. Interactions may occur in the soil or at a plant level.
Soil particles have predominantly negative charges on their surfaces. Cations adsorb to
these surfaces and remain available to plant roots. Similar cations may use the same
membrane proteins to enter the symplast. The abundance of one of them may therefore
reduce the uptake of the others. Also, movement of ions affects charge distribution across
the membrane, which in turn affects further ion movement. Optimal ratios between
nutrients in plants are therefore often as important as absolute contents. One nutrient may
replace another in some of its functions, but not in all of them.
NUTRIENT REQUIREMENTS FACTORS
- The nutrient itself (macronutrients/micronutrients)
- Plant species
- Plant age (more growth, more demand)
- Plant organ (leaves are the highest in concentration of mineral nutrients)
- Environmental conditions
NUTRIENT DEFICIENCY
Nutrient deficiency occurs when nutrient concentration in plant tissues is not enough to
support optimal growth. The causes of these nutrient deficiency can be:
- Low nutrient availability in the soil
- Poor soils (the nutrient is scarce).
 - The nutrient is present, but not available (precipitates)
- Interactions.
The consequences of nutrient deficiency are: first, a decrease in growth and yield, and
second, deficiency symptoms.
There are some differences between macronutrients and micronutrients. Toxicity is less
frequent in macronutrient than in micronutrients but may still occur. An example is the
excess of nitrogen is only tolerated by nitrophilous species.

DEFICIENCY SYMPTOMS
Foliar symptoms can arise not only from nutrients deficiencies but also from toxicity,
pathogens, or several kinds of stress. Therefore, the diagnosis of nutritional disorders by
visible symptoms could be tricky.
- Necrosis: spots of death tissue due to potassium deficiency, the symtoms appear
first in basal, old leaves, because K+ is highly mobile in the plant and it is
transported to young leaves when scarce.

- Leaf malformation: a typical symptom of Ca2+ deficiency, first shown in young

expanding leaves, because calcium moves poorly in the plant and the cell wall is
week.
 - Chlorosis: iron deficiency that have a negative effect in the chlorophyll-protein
complexes in the chloroplast and the cytochromes that participate in the electron
transport during photosynthesis. Leaf veins remains green while tissue between
veins becomes green-yellow or even yellow, in young leaves, but not in old leaves
because in the young one iron cannot be readily mobilized from older leaves to
this one. When the Fe supply restarts, the leaf returns to its normal coloration.
Under high pH conditions, a proportion of iron might be precipitated as
phosphates or other complexes in the apoplast of leaves.

Therefore, it exists chelates in fertilizers that avoid nutrient immobilization in the soil. A
chelating agent is a compound, usually organic, that can chemically combine with a metal
ion, forming a ring-like structure. Chelates are frequently used to protect metal
micronutrients (such as iron or manganese) from the chemical inactivation that so often
occurs in alkaline soils.
TOXICITY
Heavy metal in the soil (Cd, Cr, Hg, Co, Ni) are usually toxic for the plants at very low
concentration. Some micronutrients (Zn, Cu, Mn, Mo) are heavy metals.
Only some species can tolerate high concentration of heavy metals in the soil, they are
called metallophytes. These plants use different strategies of resistances:
- Exclusion of the metal, by, for example chelation by root exudate
(rhizodeposition).
- Uptake and storage, typically occurring in plants called hyperaccumulators. These
species are used to remove heavy metals from polluted soils, a process termed
phytoremediation.
Also, there are indicators which are organisms or ecological
communities strictly associated with particular environmental
conditions that its presence is indicative of the existence of these
conditions.
FOREST FERTILIZATION
The main factors to be considered to fertilize are:
- Type of fertilizer (slow-release or standard).
- Composition dosing (nutrient concentration and nutrient balances).
- Optimum time of application: age and time of the year.
- Cost (in forest practice, financial cost may become determining)
Fertilization in the nursery: most growing media can be considered infertile for all
practical purposes, therefore, nutrient loading prior to plantation in the field. Nutrient
loading in the nursery must be carefully planned, nutrients must stay stored in plant tissues
and not be used before transfer to the field.
ABSORPTION OF NUTRIENTS
Most terrestrial plants obtain both water and mineral nutrients form the soil (as inorganic
ions), but leaves are also capable of acquiring nutrients thanks to opened stomata. Nutrient
supply to the soil:
- Weathering (rock eating by mycorrhizal fungi) of parent rock material (slow
process).
- Mineralization of organic matter (recycling) is the main short- term source. They
cut the organic matter to have the minerals and change them into inorganic matter
that can be absorbed (soil microorganisms).
- From the atmosphere, mainly through nutrient deposition (with rain, snow, …)
and nitrogen fixation (except the parent rock, there is nitrogen everywhere).
The cations adsorb to the surface of soil particles, but anions tend to be repelled by the
negative charge of the surface of soil particles and remain dissolved in the soil solution
and are more susceptible of being lost by leaching. Nitrogen is a special mineral nutrient
because in can be absorbed as a cation and as an anion this is why there is a nitrogen
fixation:
The atmospheric N2 diffuses to the air gaps between soil particles. Some nitrogen-fixing
bacteria living in the soil can establish a symbiotic relationship (plat provides
carbohydrates and a home, and bacteria provides nitrogen) with the roots (nodules) of
some plant species, as in the case of Fabaceae. This N2 can be turned NH4+ in the roots
of infected plants.
The reaction is catalyzed by the enzyme nitrogenase, synthetized by nitrogen-fixing
bacteria establish a symbiotic relationship with the root of some plant species.The
industry have developed similar process to the nitrogen fixation like the Haber-Bosch
process.
Nitrogen-fixing species contribute to improve soil’s fertility, not only means of direct
nitrogen release from the roots, but also because of a particularly nitrogen-rich litter. This
improvement in soil fertility can be noticed years later, when other plant species are living
where the nitrogen-fixing species grew once.
Biogeochemical cycles:
When can start the cycle by the nutrients leached from the leaves and returned (litter),
after that there is a mineralization than consists of the breaking of the complex organic
materials and releasing the nutrients as simple ions before they become available to
plants. Then there is a nutrient absorption as inorganic molecules and simple anions and
cations. Finally, there is a translocation, the conduction of soluble material (as metabolic
products) from one part of the plant to another. These mobile nutrients can be translocated
from old to growing leaves and the essential nutrients are also translocated to the stems
before leaf shedding. Translocation in the autumn is usually better in angiosperms.
There are also some nutrients losses by leaching but are recovered thank to weathering,
the nitrogen is obtained by the N2 fixation from the atmosphere and the water comes from
rainfalls.
Nutrient recycling impairment:
Factors of nutrient recycling in forest ecosystems:
- Short-rotation forestry: energy wood plantations.
- Full tree harvest: limbs, tops and wood residue are shipped to be used as fuel
(biomass) instead of being chipped and disposed of or spread over the cut-over.
- Forest fires: nutrients lost by volatilization or/and leaching after the fire: 50 - 70%
N and other nutrients.
This is why forest fertilization should be considered when nutrient recycling is not
expected.
NUTRIENT AVAILABILITY
Soil pH is a major factor in determining the availability of nutrient elements in soils. All
essential nutrients are available in the pH range of 5.5 to 6.5. alkaline soils strongly reduce
the availability of phosphorus and iron.

Example: Chlorosis paradox is often observed in plants growing in calcareous soils,

where the Fe concentration in chlorotic leaves may be like or even higher than that in
green leaves. If there is chlorosis, but an adequate or high Fe content in leaves it can be
either to a high pH condition (a proportion of iron might be precipitated as phosphate or
other complexes in the apoplast of leaves) or other causes like low leaf ferric-chelate
reductase activity.
This nutrient availability can be change by the rhizosphere which is the immediate
microenvironment surrounding the root. Plants may change the physicochemical
properties of the rhizosphere, and thus different from those of the bulk soil.
The changes of the rhizosphere is due to rhizodeposition, release of substances from the
roots to the soil,
- Proton extrusion by roots may reduce rhizosphere pH by more than 2 units from
that in the bulk soil. Plants can also reduce rhizosphere pH by excreting organic
acids. By lowering soil pH calcicole species increase the availability of P and Fe
to the roots.
- Rhizodeposition of carbohydrates favors symbiotic associations such as those
established with nitrogen fixing microorganisms or mycorrhizae.
- In flooded soils, plant roots may even excrete oxygen. Oxygenation of the
rhizosphere leads to detoxification of potentially toxic ions that accumulate in
flooded soils.
Calcifuge species avoid calcareous soils. The lack of a high capacity to utiliza the form
of iron and other trace elements that prevail in alkaline soils may be the cause of failure
of establishment of calcifuges in such soils.
The plants can change nutrient availability in the rhizosphere thanks to the mycorrhizae,
symbiotic associations between fungi and plant roots. There are two major classes of
mycorrhizal fungi (all phosphorus and nitrogen the last one):
- Ectotrophic mycorrhizal (SEM) fungi typically form a thick sheath or mantle of
mycelium around the roots and some mycelium penetrates between the cortical
cells. They are exclusive of tree species, and they don’t enter the cells.
- Arbuscular mycorrhizal (TEM) fungi do not produce a compact mantle. The
fungal hyphae penetrate individual cortical cells, where it forms oval structures
called vesicles and branched structures called arbuscules.
- Much less abundant (mainly in the Ericaceae family), the ericoid mycorrhizal
fungi colonize the host’s epidermal root cells and do not produce a mantle.
Ericaceous plants produce polyphenol-rich leaf litter that leads to accumulation
of organize matter and recalcitrant forms of nitrogen in soils. Some ectotrophic
and ericoid mycorrhizal fungi release into the soil enzymes that provide the host
plant with an access to complex organic sources of N.
In this symbiosis, the plants give carbohydrate to fungi and the fungi gives them an
enhanced absorption of phosphorus. The association of mycorrhizae with plant roots
facilitates the uptake of immobile nutrients such as phosphorus more than that of highly
mobile nutrients. By extending around the root, the external mycelium improves
phosphorus absorption, leading to a transport rate more than 4 times higher than that of a
root not associated with mycorrhizae.
Mycorrhizal associations show low specificity, one fungus species may associate with
several plant species, and all the way around.
Nutrient movement in the soil
- Bulk/mass flow: rapid transpiration in plants may result in substantial nutrient
transport to the roots via mass/bulk flow. These movement may be jeopardized by
decreasing the water potential gradient between soil and roots (saline soils, very
dry soils) (y(soil) > y(root)).
- Diffusion: nutrient absorption by roots generates a concentration gradient between
bulk soil and root surface. It is the net movement of a substance along a
concentration gradient due to random thermal agitation. Low water availability
 reduces diffusion rates of ions below values in moist soils, because air replaces
water in pores of dry soil, greatly lengthening the path from the bulk soil to the
root surface (increased “tortuosity”).
Soils typically contain macronutrients in very dilute solutions, mineral nutrients are made
available to roots by ion movement toward roots by diffusion and by mass flow of water.
The concentrations of N, P, K often are so low in the soil solution that mass flow of water
provides only a small amount of a plant’s needs. Hence, most of these elements move to
the root surface by diffusion. Other ions are delivered more rapidly by mass flow than
they are required by the roots. Highly mobile elements move readily to the root surface.
Uptake kinetics
Nutrient uptake by roots increases in response to increasing nutrient supply up to some
maximum uptake where a plateau is reached.
- A: when nutrients are in short supply, plants tend to show a compensatory
response and the maximum uptake rate is increased.
- B: any factor that increases plant demand for nutrients appears to cause an increase
in the maximum uptake rate and conversely, at high internal concentration at the
nutrient, the capacity for uptake of that nutrient tends to be downregulated.

Compensatory
response

Solute transport across membrane barriers

Biological membranes are phospholipid bilayers in which proteins are embedded.
- Phospholipid molecules display both hydrophilic and hydrophobic proprieties
(amphipathic).
o The hydrophilic head is formed by glycerol a phosphate group and a
variable component attached to the phosphate group.
o Two fatty acids covalently linked to glycerol constitute the hydrophobic
tail of the molecule.
- There are two types of protein, the peripheral proteins and the integral proteins.
 - Steroid like cholesterol in our cells regulate membrane fluidity.
Membrane fluidity refers to how easily a component of the membrane changes position
in the membrane, and it may change due to temperature, abundance of steroids and the
composition of the hydrophobic fraction.
The extent to which a membrane permits the movement of a substance is called membrane
permeability. Biological membranes show selective permeability which means that
different molecules/ions move at different rates across the membrane. These
ions/molecules move across the membranes by diffusion, and the coefficient of diffusion
is 105 – 106 times smaller across the membranes than in the surrounding aquous solution.
This diffusion always proceeds spontaneously down a gradient of free energy or chemical
potential. Biological membranes are much more permeable to ions and polar molecules
than synthetic membranes made of pure phospholipids.
The transport across the membrane can be active (move against the gradient of chemical
potential (∆𝜇𝑗) (free energy per mol)) or passive (move down gradient). The first one is
due to the help of some proteins.
The difference in electric potential between the two sides of a biological membrane
generates a membrane potential, this is the amount of work needed to move a unit charge
from one side of the membrane to the other. This potential can be generated by either
passive or active transport or both.

Typical membrane potential across plant cell membranes ranges from -60 to -240 mV.
By convention, the negative sign implies that inside of the cell is at a lower electrical
potential than the outside.
Membrane transport classification according to whether energy is needed for the
movement or not (matters the concentration and the charge of molecule to transport
them):
- Passive transport: solutes cross membranes diffusing toward regions of lower
chemical potential:
o Diffusion
 o Facilitated diffusion (faster):
§ Proteins carrier: can run both passive and secondary active
transport, there is a conformational change.
§ Protein channel: faster, more or less selective, regulated,
inwardly/outwardly rectifying.

- Active transport
o Primary active transport: pumps: ion transport is directly coupled to
ATP hydrolysis.
o Secondary active transport: the plasma membrane electrogenic H+ -
ATPase (proton pump) contributes to keep the membrane potential
within an adequate range.

Large molecules (proteins) enter and exit the cell endocytosis and exocytosis. Very large
molecules (starch) are not able to cross the membrane.