FISIOLOGIA VEGETAL COMPLEMENTAR

(2013/2014)



MINERAL NUTRITION
and
SOLUTE TRANSPORT
Essential Nutrients, Deficiences and Plant Disorders
An essential element is defined as one that is an intrinsic component in the struture or metabolism of a
plant or whose absence cause severe abnormalities in plant growth, development or reproduction. If
plants are given these essential elements, as well as water and energy from sunlight, they can
synthesize all the components they need for normal growth. Table 5.1 lists the elements that are
considered essential for most, if not all, the higher plants. Hydrogen, carbon, and oxygen are not
considered mineral nutrients as they are obtained, primarily from water or carbon dioxide.
Essential mineral elements are
usually classified as macronutrients
or micronutrients according their
relative concentrations in plant
tissues (Table 5.1).
Essential Nutrients, Deficiences and Plant Disorders
Mineral deficiencies disrupt plant metabolism and function

An inadequate suply of an essential element results in a nutrient disorder manifested by
characteristic deficiency symptomes. For example, a particular deficiency might elicit a
specific pattern of leaf discoloration.
Some elements, such as nitrogen, phosphorus and
potassium, can readily move from leaf to leaf. Other
ones, such as boron, iron and calcium, are relatively
immobile im most pllant species (Table 5.4).

If an essential element is mobile, deficiency
symptomes tend to appear first in older leaves.

Deficiency in an immobile essential element
becomes evident, first in younger leaves.
Soil, Roots, and Microbes
Negatively charged soil particles affect the adsorption of mineral nutrients

Soil particles, both inorganic and organic, have predominantly negative charges on their
surfaces. Mineral cations such as ammonium (NH
4
+
) and potassium (K
+
) adsorb to the
negative surface charges of innorganic and organic soil particles
Mineral cations adsorbed on the suface of the soil
particles provide a nutrient reserve available to plant
roots. Mineral nutrients adsorbed in this way can be
replaced by other cations in a process known as
cation exchange ( Figure 5.6).

The degree to which a soil can adsorb and exchange
ions is termed its cation exchange capacity (CEC)
and is highly dependent of the soil type.

A soil with higher cation exchange capacity generally
has a larger reserve of mineral nutrients.

Mineral anions such as NO
3
-
and Cl
-
tend to be
repelled by the negative charge on the surface of the
soil particles, and remain dissolved in the soil solution.
Soil, Roots, and Microbes
Soil pH affects nutrient availability, soil microbes and root growth.

Root growth is generally favoured in slightly acidic soils, at pH values between 5.5 and 6.5. Fungi generally
predominates in acidic (pH below 7) soils; bacteria become more prevalent in alkaline (pH above 7) soils.

Acidity promotes the weathering of rocks that release K
+
Mg
+
, Ca
2
+
, and Mn
2+
and increases the solubility of
carbonates, sulphates and phosphates. Increasing the solubility of nutrients facilitates their availability to roots.

(1 ) CO
2
+ H
2
O H
+
+ HCO
3
-


The reaction (1) releases hydrogen ions lowering the pH of the soil and the H
+
displace K
+
Mg
+
, Ca
2
+
, and Mn
2+

from the surface of the soil particles.
Microbial decomposition produce ammonia (NH
3
) and hydrogen
sulfide (H
2
S) that can be oxidized in the soil to form the nitric acid
(HNO
3
) and sulfuric acid (H
2
SO
4
).

Excess mineral ions in the soil limit plant growth as the saline soil
induce salt stress to the non salt tolerant plants (halophytes)

Plants develop extensive root systems that may differ in form,
depending of the soil characteristics (Figure 5.7).
Soil, Roots, and Microbes
The development of the root system in both monocots and
dicots depends on the activity of the root apical meristem and
the production of lateral root meristems.

In general, three zones of activity can be defined in the apical
region of a plant root: the meristematic, elongation, and
maturation zones as it is shown in the diagram (Figure 5.9).



Different areas of root absorb different mineral ions.

The precise point of entry of minerals into the root system has
been a topic of considerable interest. Some researchers have
claimed that nutrients are absorbed only at the apical regions
of the root axes or branches. Others claim the nutrients are
absorbed over the entire root surface. Experimental evidence
supports both possibilities, depending on the plant species
and the nutrient being investigated.
Soil, Roots, and Microbes
Mycorrhizal fungi facilitate nutrient uptake by roots / Nutrients move from mycorrhizal
fungi to root cells

The uptake of mineral elements by roots may be modified by the association of mycorrhizal fungi with the
root system. The host plant provides associated mycorrhizae (singular mycorrhiza) with carbohydrates and
in return receives nutrients oe water from mycorrhizae.
There are two major classes of mycorrhizal fungi
that are important in terms of mineral nutrient
uptake by plants:
(1 ) Ectotrophic mycorrhizae
(2 ) Arbuscular mycorrhizae

Ectotrophic mycorrhizal fungi typically form a thick
sheath, or mantle, of mycellium around roots, and some
of the mycelium penetrates between the cortical cells
(Figure 5.12)

Arbuscular mycorrhizal fungi do not produce a compact
mantle of fungal mycelium around the root. Instead, the
hyphae grow in a less dense arrangement, both within
the root itself and extending outward from the root into
the surrounding soil (Figure 5.13).
Passive and Active Transport
The spontaneous movement of a substance driven by its chemical potential is termed passive transport

The movement of a substance against its chemical potential is termed active transport

The chemical potential for any solute is defined as the sum of the concentration, electric, and hydrostatic
potential. The importance of the concept of chemical potential is that it sums all the forces that may act on a molecule to
drive net transport.
In general, diffusion (passive
transport) always moves
molecules energetically downhill
from areas of high chemical
potential to areas of low chemical
potential.

Movement against a chemical-
potential gradient is indicative of
active transport (Figure 6.1)
Passive and Active Transport
Different diffusion rates for cations and anions produce diffusion potentials
When salts difuse across a membrane differentially permeable to the dissociated ions (e.g. KCl), an
electric membrane potential (voltage) develop (Figure 6.2).

The potential that develops as a result of diffusion is called DIFFUSION POTENTIAL.
As the diffusion occurs across a membrane the term membrane potential may be used, instead
diffusion potential
How does membrane potential relate to ion distribution ?

Because the membrane in the Figure 6.2 is permeable to both K
+

and Cl
-
ions, equilibrium will not be reached for either ion until
concentration gradients decrease to zero.

However, if the membrane were permeable only to K
+
, diffusion of
K
+
would carry charges across the membrane until the membrane
potential balanced the K
+
concentration gradient. In such case, the
difference of the membrane potential developed by the diffusion of
of K
+
across the membrane, at steady state, (K
+

oi
= K
+

io
) is
known as the Nernst potential for K
+
(E
K
+
= E
i
- E
o
).
Passive and Active Transport
Membrane Transport Processes

When the permeability of artificial phospholipid bilayers to ions and molecules is compared with that of
biological membranes, important similarities and differences become evident (Figure 6.6).

Biological and artificial membranes have similar permeabilities to nonpolar molecules and many small
polar molecules. On the other hand biological membranes are much more permeable to ions, to some
large polar molecules, such as sugars, and to water than artificial bilayers are.
Membrane Transport Processes

The reason for the higher permeability of biological membranes, compared to the artificial ones, is that
the biological membranes contain transport proteins that facilitate the passage of selected ions and
other molecules.

The general term of transport proteins encompasses three main categories of proteins: channels,
carriers and pumps (Figure 6.7).
Passive and Active Transport
Passive and Active Transport
Membrane Transport Processes

Channels enhance diffusion across membranes.
Channels are transmembrane proteins that function as selective pores through which molecules or
ions can diffuse across the membrane. The size of a pore and the density and nature of the surface
charges on its interior lining determine the transport specificity. Transport through channels is always
passive, and because the specificity of transport depends on pore size and electric charge more than
on selective binding channel transport is limited mainly to ions or water (Figure 6.8).
Passive and Active Transport
Membrane Transport Processes


Carriers bind and transport specific substances.

Unlike channels, carrier proteins do not have pores that extend completely across the membrane.
In the carrier the substance being transported must to initially bound to a specific site on the carrier
protein. Binding causes a conformational change in the protein, which exposes the substance to the
solution on the other side of the membrane.
Therefore , transport using carriers is highly more selective and slower than transport though chanels.
Carrier-mediated transport (unlike transport through channels) can be either passive transport or
secondary active transport. Passive transport through carriers is sometimes called facilitated difusion.


Primary active transport requires energy

Primary active transport is coupled directly to a source of energy such as ATP hydrolysis, an oxidation-
reduction reaction (as in the electron transport chain of mitochondria and chloroplast), or the absorption
of light by the carrier protein.
Membrane proteins that carry out primary active transport are called pumps. Must pumps transport
ions , such H
+
or Ca
2
+
. However, pumps belonging to the ATP-binding casset (ABC) family of
transporters can carry large organic molecules.


Passive and Active Transport
Membrane Transport Processes

Secondary active transport uses stored energy.

In plant plasma membranes, the most proeminent pumps are those for H
+
and Ca
2
+
, and the direction of
pumping is outward from the cytosol to the extracellular space. Another mechanism is needed to drive the
active uptake of mineral nutrients such as NO
3
-
, SO
4
2
-
, and PO
4
3
-
; the uptake of amino acids, peptides,
and sucrose; and the export of Na , which at high concentrations is toxic to plant cells.

The other important way that solutes are actively transported across a membrane against their gradient of
electrochemical potential is by coupling the uphill transport of one solute to the downhill transport of
another. This type of carrier-mediated cotransport is termed secondary active transport (Figure 6.10) .
Passive and Active Transport
Membrane Transport Processes

Secondary active transport uses stored energy.

There are two types of secondary active transport: symport and antiport. The example given in figure 6.10
is called symport (and the proteins involved are called symporters). Because the two substances moves in
the same direction through the membrane (Figure 6.11A) . Antiport (facilitated by proteins called
antiporters) refers to coupled transport in which the energetically downhill movement of protons drives the
active (energetically uphill) transport of a solute in the opposite direction (Figure 6.11B) .

In both types of secondary transport, the ion or solute being transported simultaneously with the protons is
moving against its gradient of electrochemical potential, so its transport is active. However, the energy
driving this transport is provided by the proton motive force rather than directly by ATP hydrolysis.
Passive and Active Transport
Membrane Transport Proteins

Numerous representative transport proteins
located in the plasma membrane and
tonoplast are illustrated in FIGURE 6.14.

Typically, transport across a biological membrane is
energized by one primary active transport system
coupled to ATP hydrolysis. The transport of one
ionic-species (for example H
+
) generates an ion
gradient and an electrochemical potential. Many
other ions or organic substances can then be
transported by a variety of secondary active
transport proteins that energize the transport of their
substrats by simultaneously carrying one or two H
+

down their energy gradient.

The protons circulate across the membrane,
outward through the primary active transport
proteins and back into the cell through the
secondary transport proteins. Most of the ion
gradients across membranes are generated and
maintained by electrochemical-potential gradients of
H
+
, which are generated by H
+
pumps.

Ion Transport in Roots
Solutes move trough both apoplast and symplast

Because all plant cells are separated by cell walls, ions can diffuse across a tissue (or be carried passively
by water flow) entirely through the cell wallspace without ever entering a living cell. This continuum of cell
walls is called the extracellular space or apoplast.
Just as the cell walls form a continuous
phase, so do the cytoplasms of neigboring
cells, collectively referred to as the
symplast. Plant cells are interconnected by
cytoplasmic bridges called plasmodesmata,
cylindric pores 20 to 60 nm in diameter with
desmotubules that is a continuation of the
endoplasmatic reticulum (Figure 6.19).
Ion Transport in Roots
Solutes move trough both apoplast and symplast

Ions cross both the symplast and apoplast.
However, in all cases, ions must enter the symplast before they can enter the stele, because of the
presence of the Casparian strip which is a suberized layer that forms rings around cells of the specialized
endodermis and effectively blocks the entry of water and solutes into the stele via the apoplast (Figure 6.20)
Xylem parenchyma cells participate in xylem
loading.

The process whereby ions exit the symplast and
enter the conducting cells of the xylem is called
xylem loading.

Xylem loading is a highly regulated process, involving
xylem parenchyma cells, which like other living plant
cells, maintain HATPas activity and a negative
membrane potential. Transporters that speciffically
function in the unloading of solutes to the tracheary
elements have been identified by electrophysiological and
genetic approaches. The plasma membranes of xylem
parenchyma cells contain proton pumps, aquaporins, and
a variety of ion channels and carriers specialized for
influx or efflux.
END