Innovation and

Reliability

Fundamentals, plant nutrition

Quality and Speed of Response

Quality and Speed

of Response

Technical Development Department

Cosmocel, S.A.
31/10/2008
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Table of Contents
PLANT NUTRITION .............................................................................................................................................. 8
Pentose Phosphate Cycle.............................................................................................................................32
Biological Work. .......................................................................................................................................33
In every event encompassing development, hormones produced in the process of biosynthesis,
play an important role, acting as a kind of messengers who control the aforementioned changes.
34
Plant Hormones........................................................................................................................................34
Water and nutrients absorption by plants. ..............................................................................................44
Water Potential, .......................................................................................................................................46
Perspiration besides being the mechanism that results in water tension of the xylem and its ascent
in the plant also enables the distribution of water and mineral nutrients absorbed by the roots
throughout the plant. ..................................................................................................................................53
Plant nutrient absorption. ...........................................................................................................................53
Transport of nutrients via Xylem ...........................................................................................................67
Transport via phloem. .............................................................................................................................71
SOIL. ....................................................................................................................................................................72
General composition of the soil.............................................................................................................78
ORGANIC MATTER ...................................................................................................................................79
PHYSICAL PROPERTIES OF THE SOIL ......................................................................................................86
CHEMICAL PROPERTIES OF THE SOIL ....................................................................................................95
Water in soil. ...............................................................................................................................................113
Water quality ..............................................................................................................................................117
Plant nutrition .............................................................................................................................................133
The nutrients ..........................................................................................................................................133
Essentiality criteria.................................................................................................................................134
Nutrient classification. ..........................................................................................................................134
Nutrients functions ................................................................................................................................140
The function of essential elements is briefly described in this section, with the understanding
that a complete description of biochemical, physiological and ecological functions would be
excessive. ................................................................................................................................................140
Keys for the identification of nutrient deficiencies...........................................................................147
1.2.- Symptoms mostly localized ..........................................................................................................148
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2.1.- The terminal bud grows normally. ..................................................................................... 148

2.2.- The terminal bud dies after new leaves are bad formed from the tip or base. .................. 149
The laws of plant nutrition. ........................................................................................................ 149
REFERENCES................................................................................................................... ................. 154

Table 1. Mean concentration of the essential elements in dry matter which are considered appropriate
for proper growth of most plants. Adapted from Marshner, 1986 and Resh, 1992 .............................. 9
Table 2 External and internal factors that regulate growth and plant development.¡Error! Marcador
no definido.
Table 3 Photosynthetic pigments found in plants, algae and bacteria. ............................................... 14
Table 4 Comparing photosynthetic models ..................................................................................... .... 18
Table 5 Factors that influence photosynthetic activity. ....................................................................... 20
Table 6 Climatic conditions for cultivation of different species. .......................................................... 43
Table 7 Approximate equivalence between photometric, radiometric and quantum units ............... 43
Table 8 Various plant water content structures, fresh weight % ......................................................... 44
Table 9 Relative consumption for different regions (kg water/kg MS) ................................................ 45
Table 10 CR for corn growth at different levels of soil fertility. ........................................................... 45
Table 11 Nutrients proportion in nutritional supply form ................................................................... 56
Table 12 Nutritional Access and Fertilizer Position. ............................................................................. 57
Table 13 Potassium and Calcium Effect in Mg absorption in barley seedlings, concentration of each
cation 0.25 me/l, Marshner (1986) ........................................................................................... ........... 62
Table 14 Relationship between nitrate supply and chlorine content in soybean leaves. Weigel et al. (1974)
................................................................................................................................................... 62 Table 15
Ion companion effect over the absorption rate of potassium and chloride in corn plants, Lügte
and Latics (1966) ........................................................................................................... ....................... 63
Table 16 Influence in the nitrogen way of supply on cation balance: anions in mustard. Kirkby (1968).
............................................................................................................................. ................................. 64
Table 17 pH Effect on the formation of some compounds. ................................................................. 65
Table 18 Primary and secondary minerals derived from weathering .................................................. 76
Table 19 Minerals and compounds derived from rocks. ...................................................................... 77
Table 20 Organic matter effects on soil ............................................................................................... 85
Table 21 Relationship between texture, source material and other properties. ................................ 88
Table 22 Ideal porosity for each texture .............................................................................................. 92
Table 23 Bulk density of various soils ................................................................................................... 92
Table 24 Bulk density and porous space .............................................................................................. 93
Table 25 Hydraulic conductivity (cm/hr). ............................................................................................. 94
Table 26 Soil behavior through time, CIANO 1996, Valle del Yaqui, Son. ............................................ 94
Table 27 Gas concentration in different environments ....................................................................... 95
Table 28 Charge specifications of certain colloids ............................................................................. 102
Table 29 Specific surface of certain clays ....................................................................................... .... 104
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Table 30 CIC of different textural classes ....................................................................................................105

Table 31 CIC clays and other materials........................................................................................................105
Table 32 Base saturation levels, % CIC ........................................................................................................106
Table 33 Base saturation per cultivation.....................................................................................................106
Table 34 Appropriate relationships between cations ...............................................................................106
Table 35 Equivalent weight of bases ...........................................................................................................107
Table 36 Materials that provide Ca and Ca + Mg, and how they modify pH .........................................108
Table 37 Fertilizers used for magnesium and potassium intake to achieve balance of bases in the soil.
108
Table 38 Relationship between the CE in saturation extract and CE in ratio 1:2, for soils of different
textures from central Mexico. ......................................................................................................................109
Table 39 Soil classification and its overall effect on crops from Ces.......................................................109
Table 40 Base saturation percentage and PSI ............................................................................................110
Table 41 Certain crop tolerance at sodium level (as specific ion) in the base exchange, PSI. ............110
Table 42 Agricultural plaster requirements to be applied to soil according to PSI, for its reduction to
5%, considering a Da = 1.2 t/m3 and Pr=30 cm. ........................................................................................111
Table 43 Soil classification based solely on the effect of sodium and salts to the ground..................111
Table 44 Classification and management of soils based on their content of salts and sodium. ........112
Table 45 Saturation moisture content.........................................................................................................114
Table 46 Moisture content at field capacity, Palacios, 1966. ..................................................................115
Table 47 Available moisture in different textures. ....................................................................................115
Table 48 Criteria and indexes classification of water irrigation. Palacios and Aceves (1970) .............117
Table 49 Water classification for salinity, Richards et al (1954) ..............................................................118
Table 50 Maximum CE due to different salts. ............................................................................................119
Table 51 Maximum solubility of some salts found in soil at different temperatures ..........................120
Table 52 Salts solubility at 20 °C ...................................................................................................................121
Table 53 Water classification, based on their salinity potential and effective salinity. .......................122
Table 54 Water classification, based on their salinity potential and soil specifications ......................122
Table 55 Cax values to calculate the RASaj .................................................................................................124
Table 56 Water classification according to CSR .........................................................................................127
Table 57 Solubility of sodium compounds at different temperatures (g/l) ...........................................127
Table 58 CaCO3 solubility at different pH ...................................................................................................127
Table 59 Classification of irrigation water according to PSP ....................................................................128
Table 60 Diagram for the Interpretation of Riverside rules .....................................................................130
Table 61 Water classification by its B level of crop tolerance. ................................................................131
Table 62 Relative crop tolerance to the presence of boron in water irrigation. ...................................131
Table 63 Classification of irrigation waters according to their content of chlorine. ............................132
Table 64 Water Effect of Na and Cl when applied by sprinkler irrigation (Mass, 1984) ......................132
Table 65 Guidelines for interpreting the water quality and its restriction degree for agricultural
irrigation...........................................................................................................................................................133
Table 66 Essential macro-nutrients to plants and their way of absorption. Bennet, 1993. ................135
Table 67 Essential micro-nutrients to plants and their way of absorption. Bennet, 1993. .................136
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Table 68 Common ionic forms in which nutrients are absorbed by plants ........................................... 136
Table 69 Nutrients classification and beneficial elements according to their chemical properties and
physiological functions (Mengel and Kirkby modified 1987). B and Mo are transition metals or
metalloids with predominance of nonmetals. Na and Co are beneficial and essential for some species.
138

Table 70 Functions that can be performed by a structural element ...................................................... 139

Table 71 Elements and its mobility .............................................................................................................. 139
Table 72 Identification key deficiencies, Uvalle B...................................................................................... 149
Table 73 Representation of the Mitscherlich equation, units Baule. ..................................................... 151

Figure 1 Phenological stages and performance components for each stage .......................................... 10
Figure 2............................................................................................................................................................... 12
Figure 3 Plant cell ............................................................................................................................................. 13
Figure 4 Chlorophyll structure ........................................................................................................................ 14
Figure 5 Internal structure of chloroplast. .................................................................................................... 15
Figure 6 Diagram of photosynthesis light reactions for the convertion of radiant energy into chemical
energy................................................................................................................................................................. 16
Figure 7 Effect of light intensity on net photosynthetic rate of a C4 corn plant and C3 barley plant. The
C4 plant is more efficient in the use of solar energy (Larcher, W., 1980) ................................................ 17
Figure 8 Sucrose formation by the binding of glucose and fructose. ....................................................... 19
Figure 9 Demand Source Ratio ....................................................................................................................... 20
Figure 10 Photosynthate distribution............................................................................................................ 21
Figure 11 Light spectrum and RFA ................................................................................................................. 22
Figure 12 Extraterrestrial radiation and daylength at different latitudes ................................................ 23
Figure 13 Relationship between photosynthesis and luminosity.............................................................. 24
Figure 14 Light point saturation at different CO2 concentrations and constant temperature of 25°C
(Hartman et al., 1987)...................................................................................................................................... 25
Figure 15 Relationship between photosynthesis and temperature ......................................................... 26
Figure 16 Relationship between corn photosynthesis and CO2 concentrations at various
environmental temperatures. Rosenberg, N. J. 1974 ................................................................................. 26
Figure 17 Photosynthesis under a different angle of light exposure. ....................................................... 27
Figure 18 Photosynthesis and Respiration.................................................................................................... 28
Figure 19 Electron transportation and groups of proteins ......................................................................... 29
Figure 20 Relationship between photosynthesis and respiration as the temperature increases. ....... 31
Figure 21 Biosynthesis process, Source: Lehninger, 1975. ......................................................................... 32
Figure 22 Graphics relating temperature, light, CO2 and photosynthesis............................................... 39
Figure 23 Effect of soil moisture on photosynthesis. .................................................................................. 40
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Figure 24 Characteristic retention water curves (g water/g soil) for various soils, depending on matric
potential Mpa....................................................................................................................................................47
Figure 25 (left) Water potential representation at different places on the path followed by water from
the soil to the atmosphere through the plant. (Right) Water potential from the xylem to the
atmosphere through the leaf mesophyll.. ....................................................................................................49

Figure 26 Opening and closing of stomata ..................................................................... ..................... 51

Figure 27 Model of apoplastic and symplastic pathways for radial ion transport through the root to
xylem. Marshner (1986). ..................................................................................................... ................. 52
Figure 28 Types of Ion absorption and low molecular weight molecules. .......................................... 55
Figure 29 Simple diffusion through channels. ..................................................................................... 56
Figure 30 Flip-plop coupled proteins .................................................................................................. 57
Figure 31 Absorption model by transporters....................................................................................... 58
Figure 32 Proton or electrogenic pumps ...................................................................................... ....... 60
Figure 33 Common antagonism ........................................................................................................... 61
Figure 34 Potassium and sodium absorption by increasing its concentration in barley roots. Marshner
(1986) .............................................................................................................................................. ..... 62
Figure 35 Ion selectivity ....................................................................................................................... 64
Figure 36 Ion external concentration .............................................................................................. .... 66
Figure 37 (left) Effect of substrate temperature and potassium concentration in plnant's dry weight
(K1=4me/l, K2=12 me/l), Chu and toop, 1975. (Right) Effect of roots temperature and P concentration
on dry weight of the cultivated tomato plant in sand. (P1 = 1 mgP/l, P2=5 mgP/l, P3=10 mgP/l). .... 67
Figure 38 Carboxyl group and phenolic group ................................................................................. .... 78
Figure 39 Proposed structure of humic acids ...................................................................................... 81
Figure 40 Cation exchange capacity of humus and different types of soil .......................................... 79
Figure 41 State of organic matter in soil (%)........................................................................................ 80
Figure 42 Process for extracting stable humus derived from leonardite ............................................ 81
Figure 43 100 Humicrop effect on the structure of soil Andosol. Dr. Victor Ordaz Chaparro, CP,
Chapingo, Mex. .............................................................................................................. ...................... 82
Figure 44 Soil classification based on depth ........................................................................................ 83
Figure 45 Triangle for texture classification ........................................................................................ 84
Figure 46 Moisturizing pattern and texture......................................................................................... 87
Figure 47 Structure and water infiltration speed ................................................................................ 86
Figure 48 ............................................................................................................................. ................. 87
Figure 49 Whitewash reaction ............................................................................................................. 93
Figure 50 Diffuse double layer diagram ............................................................................................... 97
Figure 51 Degree of base saturation and pH ....................................................................................... 98
Figure 52 Clay Structure and isomorphic substitution ........................................................................ 99
Figure 53 PH Influence on mineral colloids and organic CIC ............................................................. 100
Figure 54 Volcanic Soil Load ................................................................................................ ............... 100
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Figure 55 Load dependent on pH ................................................................................................................. 101

Figure 56 Types of water, availability and drainage characteristics ....................................................... 111
Figure 57 Salts solubility according to temperature. ................................................................................ 117
Figure 58 Nomogram for determining RAS values of irrigation water and for estimating the
corresponging values of PSI from soil which is in balance with water. Bulletin 29 FAO, (adapted from
Richards 1954)................................................................................................................................................. 120

Figure 59 Water relative reduced index of infiltration into soil affected by salinity and RAS. Bulletin 29,
FAO. .................................................................................................................................................................. 122
Figure 60 Water Irrigation classification diagram for their CE X 106 and its RAS ................................. 126
Figure 61 Summary of the most important functions of inorganic nutrients in plants, Taiz, L. And
Zeiger, E., 1988................................................................................................................................................ 143
Figure 62 Relation of synergism and antagonism between different nutrients. Penningsfeld and
Kurzmann(1975) ............................................................................................................................................. 144
Figure 63 Liebig's Law of the Minimum ...................................................................................................... 147
Figure 64 Representation of Mitscherlich Equation .................................................................................. 149
Figure 65 Nutrient sufficiency ranges and its effect on performance .................................................... 150

illustration 1: Phenological stages and components performance

..........................................................
illustration 2 Pedogenesis processes diagram ............................................................................................. 70
illustration 3 Physical and chemical weathering ……………………………………………………………………………..72
illustration 4 Soil formation diagram……………………………………………………………………………………………….73
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PLANT NUTRITION
Nutrition is a set of processes by which living organisms acquire and transform energy from
the outside. Plant nutrition studies the processes governing the availability of nutrients in
soil or substrate, its root access, absorption, transportation, assimilation and metabolic
functions of such nutrients in plants.

Given this situation, it is useful to define our main object of study and the influencing
processes mentioned above on which the plant relies. "The plant can be conceptualized as
the phenotypic expression, in space and time, of a living being’s genetic information by
means of accumulation and interaction of organic matter highly specialized from radiant
energy and inorganic simple substances. "

"The genetic information of a plant also known as genotype, in presence of an enabling

environment is capable of generating a structural and functional order process called life or
biological process, which will manifest as a phenotype whose quality of expression will
depend on the amount and quality of energy, matter and information that such
environment provides."

Plants are constituted by organs such as roots, stems, leaves, flowers which are formed by a
set of differentiated cells in tissues (meristems, parenchyma, cambium, phloem, xylem,
among others.) Each of which performs a different function. At biochemical scale it consists
of water, CO2, O2, H+, carbohydrates, proteins, fatty acids, nucleic acids, hormones,
secondary metabolites and minerals.

The growing plant would comprise up to 90% water and 10 to 20% dry matter, which at
the same time would be constituted as detailed on Table 1.
 SYMBOL AVAILABLE FORMS CONCENTRATION RELATIVE NUMBER OF ATOMS
ELEMENT
(dry weight % )

Hydrogen H H 20 6.0 60,000,000

Carbon C CO2 45.0 35,000,000
Oxygen O O2, H2O 45.0 30,000,000
Nitrogen N NO3-, NH4+ 1.5 1,000,000
Potassium K K+ 1.0 250,000
Calcium Ca Ca ++ 0.5 125,000
Magnesium Mg Mg++ 0.2 80,000
Phosphorus P H2PO4-, HPO4-- 0.2 60,000
Sulphur S SO4-- 0.1 30,000
Chlorine Cl Cl- 0.01 3,000
--- --
Boron B BO3 , B4O7 0.002 2,000
Iron Fe Fe+++, Fe++ 0.01 2,000
Manganese Mn Mn++ 0.005 1,000
Zinc Zn Zn++ 0.002 300
Copper Cu Cu++, Cu+ 0.0006 100
Molybdenum Mo MoO4- 0.00001 1
Table 1. Mean concentration of the essential elements in dry matter which are considered appropriate for proper
growth of most plants. Adapted from Marshner, 1986 and Resh, 1992.

So there are endless internal and external factors involved in order that these processes
which allow growth and development1 of a plant, are carried out optimally and can be
expressed in their entire genetic potential. In other words the appearance or condition,
under which a plant is in a given life cycle and the final performance of this, regardless it
measuring way, is the result of the interaction of genetic factors and factors that modify
them which usually are external.

The study object of plant nutrition in this case is with the modest purpose to understand which
are and how the processes and factors work interacting in the growth and development of a
plant and likewise as far as possible, whether it lies in our hands to modify them, especially the
external, to optimal ranges or values in which the best interaction is given, the desired effect is
conducive on the plant at the right time and consequently this will lead to the best possible
performance from either the total plant component or its yield component of economic
interest. Example "A water deficit can not only reduce the harvest
1GROWTH: The accumulation of living organic matter as a result of cell division and cell elongation takes place in
primary and secondary meristems.
DEVELOPMENT: Series of changes or qualitative transformations that occur in an organism during its life cycle, also
called phenological stages, are changes closely linked to production, accumulation, and action of plant hormones
which regulate such differentiation.
index but in some crops may even increase it, particularly cotton, which in early stages under
 10

suitable moisture conditions it drops its reproductive structures, produces abundant foliage
and creates fruit more belatedly, if moisture is restricted at early stages, at the onset of
flowering and mooring, the plant restricts its vegetative growth and creates and moor fruits
in lower plant knots achieving a higher harvest index". It should be mentioned that for the
proper development of a plant according to the purposes to be achieved with it, there
should exist or the optimal conditions must be given in order to properly complete each
phenological stage at the moment it happens, should this not occur there will be hardly an
opportunity to correct what in the plant’s past did not happen and probably this will not
affect biomass production as a whole, however it is possible that the yield component of
economic interest is affected.

As it can be easily guessed this is exactly the task of technical Cosmocel, the timely
identification of any external suboptimal condition that affects certain physiological
processes, and propose a solution, in terms of improving the specific condition or prepare
the plant in order that its processes are not affected or minimize such condition.

Understanding the performance as the amount of biomass produced per unit area, it is
necessary to know which are your components in their different phenological stages (Figure
1).

VEGETATIVE STAGE REPRODUCTIVE STAGE 1 REPRODUCTIVE STAGE 2

ORGAN FORMATION FRUIT OR GRAIN FILLING

SEEDING OR FLORAL FLOWERING PHYSIOLOGICAL

TRANSPLANT INITIATION MATURITY

COMPONENT 1: COMPONENT 2: COMPONENT 3:

NUMBER OF PLANTS PER POTENTIAL NUMBER OF FRUIT OR GRAIN SIZE
UNIT AREA FRUIT OR GRAINS PER PLANT AND WEIGHT

R= Nº PLANTS/m2 X Nº OF FRUITS/PLANT X WEIGHT/FRUIT

Figure 1 Phenological stages and performance components for each stage.

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Thus for each species or particular crop the performance components of interest may be
defined and focus the plantation management towards the achievement of such component.

Considered as determinants of performance:

1. Daily amount of light intercepted by the canopy

2. Efficiency with which the intercepted light is used in the production of new dry
matter.

3. Dry matter loss ratio

4. Growth duration and interest production of a plant or product

Different factors can affect one or more performance factors; such factors can be classified
into internal and external:

EXTERNAL FACTORS (ENVIRONMENTAL)

CLIMATE LIGHT, TEMPERATURE, WATER, TIME OF DAY, WIND, AND GASES (CO2, O2, N2,
SO2, nitrogen oxides, F, Cl, 03)

EDAPHIC TEXTURE, STRUCTURE, ORGANIC MATTER, POROSITY, CATION EXCHANGE

CAPACITY, PH, BASE SATURATION, AND AVAILABILITY OF NUTRIENTS

BIOLOGICAL WEEDS, INSECTS, PATHOGENS, NEMATODES, HERBIVOROUS AND SOIL

MICROORGANISMS.

INTERNAL FACTORS (GENETIC)

PHOTOSYNTHESIS ASSIMILATION PHOTOSYNTHETIC RATE

RESPIRATION ENERGY PRODUCTION, LOSS RATE OF DRY MATTER

BIOSYNTHESIS PARTITION OF ASSIMILATED AND NITROGEN: CARBOHYDRATES, PROTEINS,

LIPIDS, NUCLEIC ACIDS, ENZYMES, HORMONES.

WATER AND
NUTRIENTS
ABSORPTION AND
TRANSLOCATION

TRANSPIRATION
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GASES EXCHANGE

ENZYME-HORMONE A).- CELL DIVISION, B).- CELL GROWTH OR ELONGATION,

GENE- REGULATION
C).- CELL DIFFERENTIATION OR SPECIALIZATION

Table 2 External and internal factors that regulate growth and plant development

As mentioned before performance is the amount of biomass produced per unit area, and
we can now say that it is the result of carbon dioxide net photosynthetic assimilation or
otherwise the balance in a given time or at the end of the cycle between photosynthesis
and respiration:

Performance = photosynthesis - respiration.

Light Light saturation

High saturation High point

Light point point Light point

Producti

compensation raw photosynthesis compensation raw photosynthesis

Produ
ction
on

net photosynthesis
C

net
O

2
2

photosynthesis
Respiration
Respiration
0 0
C

sor
Ab

pti
sor
Ab

on
pti
on

CO2 Production CO2 Production

illumination intensity illumination intensity

High High

Determined relative amount of net photosynthesis by

photosynthesis and respiration
Figure 2

Photosynthesis, as synthesis process, and respiration together with the pentose phosphate
cycle as degradation processes, constitute plant metabolism, reason why we will focus on
the analysis of these processes from a nutritional point of view and not necessarily from a
scientific one.
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Photosynthesis

Photosynthesis consists in generating compounds (with lots of chemical energy stored) such as
carbohydrates from CO2 from the atmosphere and H20 with light energy derived from the sun
in a specialized organelle, which is the chloroplast present in all plant’s green parts.
Photosynthetic pigments are found in chloroplasts, which are chlorophylls and carotenoids.

-Plant eukaryotic cell Microbody Vacuole Mitochondria

Nucleus
Nucleolus
Rough

endoplasmic Cytosol reticulum

Ribosomes

Smooth
endoplasmic
reticulum
Chloroplasts

Plasma membrane
Golgi apparatus
Secretory vesicles Cell wall Figure 3 Plant Cell

The summary equation for photosynthesis is:

6CO₂+12H₂O ⟶light⟶chlorophyll⟶ C₆H₁₂0₆ + 6H₂O + 6O₂

Or else,
264g de CO₂+ 156g H₂O 686 kCal ⟶PHOTOSYNTHESIS⟶ 180g de C₆H₁₂0₆+ 78g de H₂O + 192g de O₂

For its study, photosynthesis is divided into two types of reactions:

1) Capture reactions of light energy by the pigments (Table 3) and their conversion into
chemical energy (Hill reaction). This all occurs within the thylakoid membranes present
in chloroplasts.
PIGMENT LOCATION ABSORBED LIGHT
Chlorophyll A All green plants Red blue and violet
Chlorophyll B All green plants, not found in Red blue and violet
red algae, or blue green or
diatoms.
 14

Chlorophyll C Brown algae and diatoms Red, blue and violet

Chlorophyll D Red algae Red blue and violet
Protochlorophyll Etiolated Plants Infrared and violet
blue.
Bacteriochlorophyll Purple bacteria Infrared and violet
blue.
Bacterial Viridian Green sulfur bacteria Infrared and violet
blue.
Phycocyanin Blue-green algae, red algae Orange red.
Phycoerythrins Blue-green algae, red algae Green
Carotenoids Most plants, bacteria. Blue, green-blue.
(carotenes and
xanthophylls)
Table 3 Photosynthetic pigments found in plants, algae and bacteria.

a. Capturing light energy: The uptake occurs in sensitive or photoactive

compounds at certain light wavelength known as pigments. The most important
compound of chloroplasts is chlorophyll (C55H72O5N4Mg). It has Mg at its core,
which has a great capacity for oxide reduction meaning it is able to accept and
release electrons gaining energy.

pigment carrier
membrane proteins

Figure 4 Chlorophyll structure

Higher plants synthesize chlorophyll from the simplest amino acid, glycine, and
a product of respiration, requiring light, manganese, iron, water, and the
components therein chlorophyll, C, H, O, N, and Mg.
Up to this point the importance of nutritional elements have been mentioned
(N, Mg, Mn, Fe), which they are normally applied through fertilization, and its
deficiency may be affecting the production of biomass by the affected first step
of photosynthesis, which is light capture.
 15

Ribosomes 70 S Intermembrane space

Internal Membrane
External
Membrane
circular DNA

Stroma
Cochineal
thylakoids

Cochineal
Stromal thylakoids Figure 5
Internal structure of chloroplast.

b. Conversion of light energy into chemical energy: a quite complex process that occurs in the
so-called photo system I (P 700) and photo system II (P 680), nomination given by the light
wavelength that activates (oxidizes) chlorophyll present in each PS. This takes place
through constant redox processes which the two compounds are exposed: NADP and ADP.
Managing to create a + NADPH reserve (nicotinamide adenine dinucleotide phosphate) and
ATP (adenosine triphosphate, ADP + Pi) thus accumulating chemical energy that was
initially radiating.
This step also provides the photolysis of water that releases used electrons in the PS II,
allowing the continuity of PSI and PSII, moreover its protons (H+) provide the necessary
energy for ATP synthesis, besides they are the main source of hydrogen required to convert
CO2 into glucose. Photolysis is performed involving an enzyme where Mn, Cl and Ca
participate. Nitrogen once again is involved as part of the proteins that are composed of
thylakoid compounds and enzymes, as well as, in ATP and NADP, and another integrates as
an important part of energy storage, P.
Once again we find nutrients, which normally are applied through fertilization and its
deficiency, can lead to a low conversion from radiating energy into chemical.
 16

Figure 6 Diagram of
photosynthesis
light reactions for the
convertion of radiant energy into chemical energy.

2) The enzymatic reactions where CO2 is fixed and reduced involving chemical energy
obtained in the preceding processes, ATP and NADPH forming carbohydrates (Calvin cycle).
These reactions take place in the chloroplast stroma, and are conducted by enzymes, being
one of the main the one that fixes CO2 initially and that is called ribulose bisphosphate
carboxylase-oxygenase (RUBISCO). Should be noted that such reactions occur in presence
of water, therefore a water shortage would limit the plant’s photosynthetic activity. As this
stage apparently does not require light energy, it is also referred to as the dark reactions
stage although it actually occur during the day. However, it is known that light is needed to
activate enzymes such as RUBISCO, which by the way it is the most abundant protein on
leaves whereby takes high significance maintaining appropriate levels of the elements that
conforms it and even the external application of L-amino acids in order to maintain their
synthesis at all times.
The overall equation for this stage can be represented as,
6CO₂+ 18ATP + 12 NADPH⁺ ⟶ C₆H₁₂0₆ + 18ADP + 18Pi + 192 g de O₂

However, three paths are known for reducing CO₂ into glucose and which itself defines
three groups of plants: C3, C4 and MAC, because of the first stable compound that is
formed with the added CO2, at C3 is three carbons and C4 4 carbons; in either case the CO2
used is obtained by the plant during daytime, which penetrates to the chloroplast stroma
after overcoming several resistance: boundary layer, stoma, stomatal camera, cell
membrane and chloroplast membrane. In the case of MAC plants reducing CO 2 into glucose
is given by a path similar to C3, with the exception that the CO2 it uses comes from an
organic compound formed at night when these plants have access to it by having their
stomata open only then as a protection against water loss.
 17

Barley, C3
Plant, low
photosyntetic
capacity.
Corn, C4
Net Photosynthesis

plant, high
photosyntetic
capacity

Ligh intensity

Figure 7 Effect of light intensity on net photosynthetic rate of a C4 corn plant and a C3 barley plant. The C4 plant is
more efficient in the use of solar energy (Larcher, W., 1980)

FEATURE C3 PLANTS C4 PLANTS MAC PLANTS

Wheat, barley, Corn, sorghum, Pineapple, agave,

potatoes, beans, rice, sugarcane, millet, cactus
rye, tomato bermudagrass

% Of the world's Flora in 89% <1% 10%

No. of species.

Typical habitat Pattern undefined Warm regions and Xeric and epiphytic
grasslands. regions.

Leaf anatomy Palisade cells + spongy There is no cell There is no cell

parenchyma differentiation in the differentiation in the
mesophyll, but it has a mesophyll, but it has large
great bunch of sheath cells cells with large vacuoles.
(Kranz anatomy)

Light saturation point 6000 footcandles 8000 y 10000 6000 footcandles

footcandles

First stable product of PGA Malate Malate

CO2 fixation

Photorespiration Up to 40% of Undetectable Undetectable

photosynthesis

Compensation point 40 a 100 ppm 0 a 10 ppm 0 a 10 ppm (night)

(CO2)
 18

Intracellular CO2 in 200 100 10000

daylight

Stomatal frequency 40 a 300 100 a 160 1a8

(stomata / mm2)

EUA (g of fixed CO2 per 1 –3 2–5 10 – 40

kg of transpired H2O)

Maximum growth rate 5 – 20 40 – 50 0.2

(gm-2d-1)
Table 4 Comparison of photosynthetic models

Performing a quick analysis we may observe that the least efficient plants in the production
of dry matter and in relation to water usage are the C3 plants representing the highest
percentage in number of species. It is possible to notice that if a MAC plant will improves
especially its moisture conditions, it may have a high productivity by using its
photosynthetic advantages.

Photorespiration

Along with photosynthesis, during daytime (in presence of light) another metabolic process
called photorespiration is conducted, which is also initiated in the chloroplast and through
the same enzyme responsible for setting CO2 into photosynthesis (RUBISCO). In this process
two other organelles participate, mitochondria and peroxisome. It seems that one of the
main functions of this process "in vivo" is the removal of excess reducting power (energy)
and keep oxygen busy when stomata are closed and there is a CO2 deficiency inside the cell.
This may occur when there are conditions of high light and temperature intensity and
deficiencies in water supply. That is to say, as a result of high temperatures the
photosynthetic efficiency notoriously decreases since instead of attaching CO2 it detaches
it, a formed amino acid is also deaminated, therefore the released NH3 will require energy
to rejoin the metabolism; increasing the temperature will decrease proportionally the
solubility of CO2 more than that of O2, on the other hand the increase in lightness will
directly cause the release of O2 in chloroplasts of H2O and the CO2/O2 ratio will decrease,
this implies an increase in the photorespiration rate.
 19

As it may be noted the decrease in photosynthetic rate may occur for various circumstances
which act in an inter-related manner, it is important being able to interpret and take measures
as in the latter wherein it is affected by high temperature and luminosity bringing as a result an
amino acid degradation, where the plant will have to synthesize again such amino acid
requiring energy for this purpose or another amino acid that may be caught externally allowing
the plant, either to use energy for carring out protein synthesis and not the synthesis of such
amino acid, or otherwise if the amino acid in question is not provided and the plant must
synthesize it, if other amino acids would be provided thus the plant would no longer need to
synthesize them momentarily thereby saving energy, the objective would a positive energy
balance for the plant before a stress situation.

Most important Photosintates:

a. Carbohydrate reserves. High molecular weighte and not transported in the plant.
i. Starch, formed by glucose.
ii. Inulin, formed by fructose.
b. Carbohydrate transport. Low molecular weight and non-reducing.
i. Sucrose. It is the most abundant and can be found in the phloem.
ii. Raffinose.
iii. Stachyose.
iv. Alcohol Sugars. Sorbitol y mannitol.
c. Carbohydrates such as glucose of immediate incorporation into structural
polysaccharides such as cellulose and hemicellulose of the cell wall, or else as
substrates entering the respiration process.
GLUCOSE
FRUCTOSE

SUCROSE

Figure 8 Sucrose formation by the binding of glucose and fructose

 20

Factors that influence photosynthetic activity.

INTERNAL AND MORPHOLOGICAL EXTERNAL

Pigments Light energy
Enzymes Carbon dioxide
Stomata Water
Age and leaf number Temperature
Demand source relations Nutrition
Plant shape
Leaf area index
Seeding density
Seeding pattern
Table 5 factors influencing photosynthetic activity.

Within the internal factors one of the most important from a practical standpoint, since it is
something that can be handle agronomically, is that of source-demand relations, being the
basis for practices such as cropping, defoliation, flowers or fruits thinning, pruning.

High Radiation
Large MeanT
Source Not limited Water and
nutrients
Source
Demand
Many Young fruits
Few assimilated Many assimilated
consumed by fruits (Low consumed by fruits (Low
charge) charge)

Few Young fruits

Source
Demand

Low Radiation Small

MeanT Source
Limited Water and nutrients

Figure 9 Demand Source Relations

The demands can be classified into different categories as described below:

 21

a. Consumption: meristems and young organs that consume soluble sugars and is a
short-term demand.
b. Accumulation: stems, roots, fruits, seeds where long term soluble sugars and starch
accumulate.
c. Secretion: Nectaries, also a short term demand with soluble sugars.

Each plant species channels its photosintates to different parts of the plant in
different proportions according to their priorities.

Figure 10 Photosintates distribution.

As for the external factors it can be mentioned in a summarized form that light energy is
one of the most important factors constantly acting in short (daily) and long term (weeks,
months) over the control rate of photosynthesis. The daily amount of light, its average
intensity and quality have a great influence on this process and consequently on the
performance thereof. The amount refers to day length regarding the night, photoperiod,
and this related to latitude and time of year; the intensity is the amount of photons that
affects space per unit and time; and quality refers to the different wavelengths of the
spectrum that are incident on a particular location, a greater excitation of chlorophyll can
be achieved in the range between 430 and 480 nm and between 595 and 620 nm. Within
this it is worth mentioning another concept normally used, photosynthetically active
radiation (PAR), whose wavelength ranges from 400 and 700 nm, ie, the spectrum of visible
light that influences the photosynthetic process. It is estimated that approximately
between 40% and 50% of the total radiation is PAR.
 22

Wave Name Hard X rays Ultraviolet Infrared Radio Waves

Gamma Rays Smooth X rays Microwaves

Radioactive X rays Sun People Microwave

elements

Transmitters

Greenish Bluish Yellowish

Violet Blue blue Green Green Green Yellow Orange Red

Wavelenght

Figure 1 Light spectrum and PAR

 23

Extraterrestrial Radiation annual assessment (Ra) in the Equator, 20º and

40º north and south latitude

Equator
Day -1

Jan Apr May Aug Dec

Daytime light hours annual assessment (N) in the Equator, 20º and 40º north and south
latitude
N maximum sunlight (Hours/day)

Equator

Jan Apr May Aug Dec

Figure 12 Extraterrestrial radiation and daylength at different latitudes.

 24

LUMINOSITY-PHOTOSYNTHESIS ENERGY RATIO

Plant

Plant Sun

Net Photosynthesis

Plant Shadow
Respiration

Figure 13 Relationship between photosynthesis and luminosity

It is observed in the previous chart the light compensation point where a certain amount of
light would be setting the same amount of CO2 through photosynthesis than the one it would
be producing by the effect of respiration; as the luminosity increases the light saturation point
will appear, which represents the maximum capacity of such group of plants to fix CO2 at a
given temperature and a CO2 concentration if temperature changes up to a certain limit and if
the concentration of CO2 rises, the saturation light point will increase as well.
 25

Being CO2 the base of organic compounds is very important likewise, due to its low
concentration at which it exists in the atmosphere, is generally a limitation to
photosynthesis, given the photosynthetic response which most of the plants have to a
higher concentration of CO2.
Relative rate of photosynthesis

LIGHT INTENSITY (lux X 10 3 )

Figure 14 Light saturation point at different CO2 concentrations and constant temperature of 25°C (Hartman et al., 1987).

When the concentration of CO2 increases, photosynthetic activity rises. Note that the light
saturation point increases when increasing the CO2 concentration.

Temperature is another important external factors affecting the activity of enzymes and
indirectly affects the relative humidity influencing the opening and closing of stomata. This
affects different ranges for each of the plant groups, in the case of C3 the optimal is above
25ºC near to 28°C from which the photosynthetic rate begins to diminish and accordingly
decreases performance.
 26

TEMPERATURE-PHOTOSYNTHESIS RATIO

Photosynthesis
Photosynthesis
with high supply
of CO2

Photosynthesis

Light Respiration
with natural
supply of CO2

Temperature
Figure 15 Ratio between temperature-photosynthesis
PHOTOSYNTHESIS

Figure 16 Ratio between corn photosynthesis and CO concentrations at different environmental

temperatures. Rosenberg, N. J. 1974
 27

The concentration of P in cells is critical in photosynthetic efficiency, as noted in the chart

below, as there is a greater concentration of P, CO2 fixation is performed efficiently even in
low CO2 concentrations. Moreover, it is said that when phosphorus is deficient for growth,
sugar distribution changes in the plant and starch synthesized in leaves changes its
features. This starch has less associated phosphate, less amylopectin and is less susceptible
to β-amylase degradation; translating this into an inefficient use of carbohydrates for its
traslocation into areas of demand.
CO2-PHOTOSYNTHESIS AND P RATIO
Net Photosynthesis, A

Intracellular CO2 Concentration

Figure 17 Ratio between photosynthesis, CO2 concentration and P concentration

PAR Importance
With PAR deficiencies, ATP and NADPH+ formation decreases, CO2 conversion into CH2O
decreases, therefore there would be fewer products for cellular respiration and
biosynthesis and as a result there would be less growth and performance. Changes occur in
leaves’s cellular differentiation (thinner leaves with fewer chloroplasts and less
photosynthesis capacity with high radiation, elongation or etiolation (auxinic syndrome),
and nutritional deficiencies of P, K, Fe, and N are presented.

On the other hand PAR excess turns up with free radicals and protective substances
(carotenoids, violaxanthin) occurance, the cell membrane on upper leaves is damaged with
eventual chlorophyll destruction (photo-oxidation), scrolling leaves by starch excess, will
lead to changes in cell differentiation of leaves making leaf cuticles thicker, more
chloroplasts and greater photosynthetic capacity with high radiation; damages may occur
to fruits (burns, maturation des uniform).
 28

Is possible to perform various practices to make effective the use of light by plants to
achieve higher conversion into chemical energy:

 Vegetable patch or crop designing in order to possess adequate spacing and

 orientation.
  Plant designing by pruning.
  Achieve better development and foliar area duration
 Adequate distribution of dry matter towards the interest portion
o Reproductive or vegetative competition
o Leaves into fruits relationship
CORN SHAMROCK
Sunlight
Leaf Photosynthesis

(Total sun light %)

Leaf Photosynthesis for Corn (M) and Shamrock Simple models and calculations of
(T) as function of ilumination (l). These species photosynthesis population por corn and
ilustrate different saturation levels at solar Shamrock with two foliar distributions: con a =
exposure. 0º, ilumination (l) 90% and 10% total sun light;
with a = 60º, l 50% and 50% of total sunlight.

Figure 18 photosynthesis under different angle of light exposure.

Respiration
It is the main process for carbohydrates degradation in order to liberate stored chemical energy
and use it in the processes of growth, differentiation, absorption and transport of compounds.
Respiration occurs in the mitochondria and is through controlled oxidation of a part of these
carbohydrates to capture and store a portion of the released energy in the form of ATP and the
reducing power primarily as NADH+. Enzymes achieve control. Being a gradual process will also
generate various intermediaries that may exit this process and therefore be used as precursors
to form lipids, amino acids, nitrogenous bases, pigments, etc.
 29

Figure 19 Photosynthesis and Respiration

The equation that summarizes respiration is usually expressed as follows:

C₆H₁₂0₆ + 6O₂ ⟶ 6CO₂ + 6H2O + Energy

Or else:
C₆H₁₂0₆+6O₂+36ADP + 36Pi⟶ 6CO₂+6H2O + 36ATP,
∆G = −686 kcal/Mol

Note the amount of liberated energy per molecule of glucose and the essential involvement
P once again in this process.

For its study, respiration is divided into three phases or stages::

1) Glycolysis. It is a metabolic process of anaerobic character that takes place in the

cytoplasm. In this process a partial breakdown of carbohydrates takes place, releasing
chemical energy. Glucose is converted into two molecules of pyruvic acid.
Glucose + 2 NAD + 2ADP +2 Pi → 2 Pyruvic Acid +2 NADH+H+ + 2ATP

2) Krebs cycle. Wherein the pyruvic acid is oxidized to CO2 releasing a greater amount of
chemical energy. Eliminating the H gradually and adding itself to compound electron
transporters, present in the inner membrane of mitochondria, will release pyruvic acid
energy. The main result is the formation of a reducing power reserve in the form of
NADH+ and amin-flavin dinucleotide (FADH2), carried out in the mitochondrial matrix as
an aerobic process.
 30

2 Pyruvic Acid+8NAD+2FAD+2ADP+2Pi →6CO2 + 8NADH+H +2FADH2 +2ATP

3) Electron transport and ATP formation (oxidative phosphorylation). During this phase the
NADH+H+ from glycolysis and Krebs cycle plus the FADH2 from Krebs cycle as well enter
electron transportation in order to obtain ATP. O2 is consumed at this stage, which is
reduced to H2O. It is held in the inner membranes (crests) of mitochondria. And can be
summarized as follows:
2NADH+H+ + 8NADH+H+ + 2FADH2 +36ADP + 36Pi + O2 →
10 NAD+2 FAD+36ATP+ H2O

Figure 20 Electron transport and group of proteins.

The electron transporters in the mitochondrial membrane are groups of proteins,

embedded as well in the inner membrane of mitochondria ATP synthase can be found
synthesizing ATP from ADP and Pi.

Resulting thereof is clearly noted the importance of participation by elements such as N, P,

Ca and O2 in the respiration process.

There are different inhibitors of this last phase of respiration with different action
mechanisms. These compounds may be some poisons, antibiotics, compounds used as bud
 31

burst stimulators and delaying senescence. Examples: antimicin, cyanide, carbon monoxide,
dinitrophenol, oligomicin.

Certain aspects noteworthy about the respiratory process:

 Respiration takes place throughout the plant unlike photosynthesis, which is only
performed on green areas. In a mole of glucose containing ∆G = +686 kcal, 36 moles
of ATP are synthesized, each storing approximately 7.3 kcal for a total of 263 kcal,
this represents a process efficiency of 38%.

 Molecular oxygen plays an important role as final electron acceptor and oxidizing
agent.

 As a result of the process a fraction of glucose is transferred to ATP. This compound
and to a much lesser extent, is responsible for transporting energy to those
processes or cellular reactions that require it. That is to say ATP energy is used for
 biological work.
 The main factors affecting respiration are:
o Temperature
o Presence of water
o Amount of breathable material
o Presence and oxygen uptake by the roots and
stomata o Amount of CO2
o Type of cell wherein is performed
 32

Optimum temperature range for *Net photosynthesis is the difference

between raw photosynthesis and respiration
the net photosynthesis
Respiration
Net photosynthesis Respiration
Net gain from Net loss from
carbohydrate production carbohydrate production

Temperature ºC
Critical temperature in which the amount of
CO2 consumed in photosynthesis is equal to
the amount released in respiration

Figure 21 Relationship between photosynthesis and respiration as the temperature increases.

Regarding oxygen is noteworthy saying despite being metabolically important when

"disengaged", it can cause in plant serious oxidation problems being able to induce the
formation of reactive oxygen species (ROS) such as hydrogen peroxide and free radicals as
superoxide. The plant has different mechanisms to degrade ROS and eliminate free radicals.
Otherwise these compounds lead to alterations in the genetic material, membranes and
enzymatic processes leading to rapid aging and eventually cell death.

Pentose Phosphate Cycle

It is a carbohydrate degradation process conducted in hand of glycolysis in the cytoplasm
and is anaerobic. The main purpose of this cycle is to produce intermediates different of
those occurring in respiration, as well as generating a different form of chemical energy,
which is the NADPH +H+. Among the most important intermediates produced in this cycle
are: Ribose-5 P, an essential part of the nucleotides and erythrose -4 P, which, together
with phosphoenol pyruvate formed in glycolysis participate in the synthesis of a large
number of compounds in the shikimic acid path, within the so-called "secondary
metabolism", generating phenols, alkaloids, lignin, amino acids, etc..
 33

Biological Work.
Is the use of chemical energy to perform the processes that sustain life:

1. Biosynthesis work:
a. From simple precursors such as H2O, CO2, NH4, SO4--, H2PO4-, K +, Mg ++,
Ca++, etc. in presence of enzymes, the most simple organic molecules such
as amino acids, nucleotides, monosaccharides, fatty acids, and other organic
acids are synthesizedto, to which the name of basic structural units is given.
b. These are chemically bonded to form macromolecules such as RNA and DNA,
proteins, polysaccharides, lipids and hormones.
c. With the aggregation, interaction, and interdependence of macromolecules,
supra molecular systems such as membranes, ribosomes and enzyme
systems are integrated.
d. Subsequently cellular subsystems such as mitochondria, the endoplasmic
reticulum the nucleus, chloroplast, etc. are formed.
e. Finally the interaction and interdependence of cellular subsystems results in
the formation of cells, thereafter these shall form tissues, organs, etc. which
translates into growth and development.

SIMPLE BASIC ORGANIC

MOLECULES MACROMOLECULES
PRECURSOR

CELLS ORGANELLES SUPRAMOLECULAR

SISTEMS

Figure 22 Biosynthetic process, Source: Lehninger, 1975

The order and sequence of these processes occur determined by the genetic
information in each cell. This all happens using ATP energy and the reducing power of
the compounds NADPH+, NADH+ and FADH2. Former solar energy. In this kind of work
most of the energy is used from respiration.

2. Concentration or Osmotic work: Energy expenditure carried out in cells for

transporting and concentrating substances even against the laws of diffusion,
 34

these substances can be nutrients or other types. The movements can be

absorption or excretion.

Active transport is the name given to the active movement of solute molecules
against their tendency to disperse (diffuse) and involves an expenditure of energy.
This action of “pumping” maintains the cell relatively constant and maintains an
optimal internal environment for life, even though the external environment might
have a different chemical composition.
3. Mechanical work: Refers to energy expenditure by the cell to perform structure
movements, ie separation of chromosomes in meiosis and mitosis, cytoplasmic
movements, among others.
4. Data collection. Data collection in cells is considered a type of energy.

The continuity and quality of biological work are responsible for growth, reproduction, cell
differentiation and organization, therefore the production of a plant, and they depend
mainly on the energy captured and transformed by photosynthesis and re-transformed by
respiration.

Development
Processes of growth and differentiation are carried out as a result of biological work, which
are the basis for development, which comprises a series of controlled events, connected
and integrated by the genome, leading to initiate and complete a plant’s lifecycle. The
handling that is given to the plant and the environmental conditions under which they
occur modifies these events.

Development encompasses both cell division and cell expansion and is conducted in the
primary and secondary meristem. Cell differentiation involves changes in enzymes leading
to changes in cell function generating different tissues (xylem, phloem, parenchyma,
epidermis, etc.) and organs (roots. stem, leaves, flowers, fruits, seeds) as a result.

In every event encompassing development, hormones produced in the process of

biosynthesis, play an important role, acting as a kind of messengers who control the
aforementioned changes.

Plant Hormones
Are organic compounds synthesized at low concentrations in the plant, in a particular area,
which act at the place of synthesis or may be transported great distances to exert their action.
A compound mechanism of action is very important to consider it as hormone. Hormones can
act by inducing or repressing the synthesis of any enzyme, or either activating or inhibiting any
enzyme that may already be present, and in some cases the hormones can
 35

alter the permeability of the cell. Is worth mentioning that in order to control a process,
hormones act based on its balance with one another. The main plant hormones are:

1. Auxin (indole acetic acid), its precursor the L-tryptophan.

a. Synthesis in meristems
b. Transport via phloem parenchyma, cortex and medulla.
c. Cell division and development
d. Differentiation of the vascular system
e. Induce apical dominance
f. Delay senescence
g. Responsible for tropisms
h. Induce rooting cuttings and seedlings
i. Induces fruit mooring, especially in fruit species with many seeds.
j. It is an active ingredient of some herbicides.
2. Gibberellins. Its precursor is mevalonic acid, acetyl-CoA
a. Synthesis in younger leaves roots and seeds.
b. Transport via xylem, phloem, bark and marrow
c. Inhibits lethargy or removes this from plants and promotes seed germination
and sprouting.
d. Increase length internodes, keeping the number of internodes.
e. Induces flowering in long day and vernalization plants.
f. Stimulates pollen tube growth.
g. Promotes synthesis of amylases.
3. Cytokinins. Its precursor is adenine (glycine, CO2).
a. Young fruits synthesis, seeds, young leaves and roots.
b. Transport via xylem.
c. It stimulates cell division.
d. Induces chloroplast formation
e. Promotes organogenesis differentiation and activation of flower buds.
f. Stimulates leaf expansion.
4. Abscisic acid. Its precursor is mevalonic acid, acetyl CoA.
a. It is synthesized in chloroplasts, leaves, stems and fruits. It is present in seeds.
b. Extends latency of buds.
c. Extends the seeds rest.
d. Inhibits the germination of seeds that have completed their rest.
e. Inhibits the transport of IAA
f. Responsible for senescence and leaves death.
g. Under water stress promotes stomatal closure and provides resistance to
oppression.
 36

h. Stimulates the transport in xylem

i. Transport via xylem and phloem.
5. Ethylene. Its precursor is methionine and ACC (amino-cyclopropane carboxylic acid)
a. Is synthesized in any area of the plant.
b. Causes stems and petioles epinasty
c. Inhibit the elongation of stems roots and leaves, to inhibit auxin transport.
d. Promotes leaf senescence. In balance with cytokinins.
e. Promotes abscission of leaves and fruits. In balance with auxin.
f. Causes stomatal closure and accelerate senescence.
g. Stimulates the production of aerenchyma and latex formation.
h. Inhibits root development and promotes the production of adventitious roots.
i. Stem basal thickness increases. Inducing lateral cell expansion.
j. Accelerates and standardizes climacteric fruits ripening.
k. Promotes flowering in pineapple.
Recently there have also been incorporated into the brassinolides.
6. Brassinolides.
a. Present in all plant tissues.
b. Promote elongation and cell division in stem segments.
c. Promote leaves unwinding.
d. Promote thedevelopment of pollen tubes.
e. Promote xylem differentiation.
f. Promote germination.
g. Inhibit the growth of roots.
h. Delay the abscission of foliage
There are other compounds that also synthesize and control plant development
although it is still unclear whether they act as hormones: Phenols, polyamines,
jasmonates.

Considerations on hormones.

The primary action of a hormone in a specific process depends on: a) synthesis rate, b)
transport, c) degradation rate, d) the way is found, active (unconjugated or inactive), e)
presence of specific receptors, f) concentration of other hormones.

Effect of the environment on hormones.

1. Gravity promotes lateral auxin transport in stems.

2. Gravity stimulates lateral distribution of ABA in roots.
 37

3. Heat stimulates auxin transport in the stem and inhibits FA and cytokinins synthesis.
4. Foliar wilting indicates ethylene and ABA production.
5. The cold induces ethylene sheet, in gibberellic acid and cytokinins seeds, causing
reduction in ABA seed.
6. The flooding promotes ACC accumulation (ethylene precursor) in xylem of stems
and roots, and synthesis of ABA in roots.
7. The water deficit stimulates ABA synthesis and inhibits kinetin (cytokinin) in the root
apex.

There are a lot of natural or synthetic compounds, which can be applied to similar plants or
chemically different, they are called "hormonal type growth regulators". Used in plant
production to induce or inhibit a specific process. These regulators may act by modifying
the concentration, synthesis, transportation, degradation or any hormone receptor.

As noted above the action of a hormone or onset of action is determined by many factors
including light as one of the most important.

Light effect on growth and development.

The light acts indirectly through photosynthesis and directly in the process of photo
morphogenesis, by which plants capture light at different wavelengths and these luminous
signals generate physiological changes that affect growth, development and plant
differentiation. The involved molecules are:

  Phytochrome, capture blue and red light (660 to 730 nm)

  Cryptochromes, capture blue light between 350 and 366 nm, at 450 nm
  UV photo receivers they absorb UV radiation between 280 and 320 nm
 Photochlorophylline, absorbs red and blue light, as reduced it becomes chlorophyll A

They are called photoreceptors and nutrients such as P and Ca are involved its mechanism
of action.

Cryptochromes and phytochrome functions, which have a similar activity, act by regulating
the plants life cycle, seed germination, development outbreaks, flowering and senescence.

1. Photoperiodic morphogenetic answers.

a. Initiation and disruption of winter dormancy of perennials and woody plants.
b. Germination of certain types of seeds.
c. Bulbs formation in certain plants
2. Non-photoperiodic morphogenetic answers.
a. Leaves unwinding in monocotyledonous
 38

b. Stem elongation inhibition

c. Modification of geotropism and phototropism
3. Photo answers non-morphogenetic.
a. Synthesis stimulation of enzymes and proteins in illuminated plants
b. Synthesis of anthocyanins and other flavonoids
c. Gibberellins Biosynthesis.

Regarding hormonal activity the light,

a. Stimulates lateral auxin transport in the stem

b. Promotes polar auxin transport in the root
c. High light intensity generates destruction of auxin
d. Promotes the synthesis of AG in long-day and short-day plants
e. In short-day plants promotes accumulation of ABA for some species.
f. Red light promotes FA synthesis and increases the level of kinetin.
g. Inhibits polar auxin transport in the stem
h. Red light inhibits the production of ethylene in seedlings achieving standing erect.
i. Inhibits cytokine production in short-day plants.

As discussed above and now confirmed, the production of a crop is the final result of the
continuous interaction between genotype and the environment throughout its cycle. These
interactions determine the efficiency of their physiological processes or functioning
(photosynthesis, respiration and biological work) and its efficiency depends on
performance and quality.

Therefore affecting the physiological processes of the plant will affect its performance, and
these physiological processes will be affected if the plant do not have the quality, quantity
and intensity of adequate light at the place where is being cultivated, if CO 2 and O2
availability are not sufficient, if the air and soil temperature are not in the appropriate
range for cultivation and, whether the availability of water and mineral nutrients are not
sufficient, or whether they are affected by any pest or disease to such an extent that
physiological processes are inhibited.
 39

Graphics showing some the foregoing interactions are shown below.

Response rate of a leaf photosynthesis to Response rate of a leaf photosynthesis to
light intensity light intensity
Photosynthesis (%)

Photosynthesis (%)
Solar Radiation (foot-candle) Solar Radiation (foot-candle)

DIFFERENT LEAVES (ALFALFA) PHOTOSYNTHESIS RATE

A CUCUMBER LEAF PHOTOSYNTHESIS IN
Bottom Leaf DIFFERENT CONDITIONS (GAASTRA, 1959)
Intermediate Leaf
Superior Leaf
Photosynthesis

Photosynthesis

Foot-candle
 40

THE EFFECT OF CO2 PHOTOSYNTHESIS IN TOMATOES LEAVES AT

EFFECT OF LIGHT AND CO2 ON PHOTOSYNTHESIS
DIFFERENT INTENSITIES (GAASTRA, 1959)
RATE OF A C3 LEAF (AT 25 DEGREES)
Foot-candle

Relative Rate of Photosynthesis

Foot-candle
Foot-candle
Photosynthesis

Foot-candle
CO2 Concentration

LEAF PHOTOSYNTHESIS DEVELOPED AT DIFFERENT LIGHT

EFFECT OF CO2, LIGHT AND TEMPERATURE IN TOMATO
INTENSITIES (PICKEN et al, 1986)
LEAVES PHOTOSYNTHESIS (GAASTRA, 1959)
Leaf
Leaf
Leaf
Photosynthesis

Photosynthesis

Intensity in Foot-candle
 41

THE EFFECT OF CO2 PHOTOSYNTHESIS IN LEAVES OF LEAF NET PHOTOSYNTHESIS OF 4 TOMATO CULTIVARS IN
TOMATOES AT DIFFERENT INTENSITIES (GAASTRA, 1959) RESPONSE TO CO2
Foot-candle
Foot-candle
Foot-candle
Photosynthesis

Photosynthesis
CO2 CONCENTRATION
CO2 CONCENTRATION

EFFECT OF LIGHT AND CO2 ON PHOTOSYNTHESIS OF A C3

(BEAN) AND C4 (CORN) PLANT
Bean
Bean
Corn
Photosynthesis

Compensation
Point

Figure 23 Graphics relating temperature, luminosity, CO2 and photosynthesis.

 42

Effect of soil moisture on photosynthesis. Chang, 1974

Effect of soil moisture on photosynthesis. Chang, 1974

Figure 24 Effect of soil moisture on photosynthesis.

 43

TABLE OF CLIMATIC CONDITIONS FOR DIFFERENT SPECIES CULTIVATION

Minimum and optimal thermal environment level, optimal CO2 levels, relative humidity (rh) and soil temperature for cultivation of
vegetables and flowers

SPECIES Min. Letal Biological Opt. Day Opt. Day Biological Min. Opt. Soil Opt. CO2 (in
Temp. Min. Temp. Temp. Temp. Max. Temp. Germination Germination Temp ºC ppm)
Temp. Temp.
Vegetables
Cucumber
Melon
Zucchini
Beans
Pepper
Aubergine
Lettuce
Tomato
Flowers
Carnation
Rose
Gerbera
Chrysanthemum
Hydrangea
Tulip
Iris
Narcissus
Cyclamen

Tables obtained from the Book '' MODERNE TECNICHE DI PROTEZIONE IN HORTICULTURE, FLORICULTURA E FRUTUCULTURA '' Romano Tesi.

Table 6 Climatic conditions for different species cultivation.

UNIT EQUIVALENT TO

1 foot candle 10. 76 lux

1 klux 4 Watt/m2
1 lux 0.093 foot candle
1 foot candle 0.1 Watt/m2 RST (total solar radiation)
1 foot candle 0.0432 Watt/m2 PAR (photosynthetically active radiation)
1 Watt 1 J/s
1 Watt/m2 1 J/s/m2
1 Watt/m2 23.234 foot candle
2
1 Watt/m 14.3 cal/min/m2
1 Watt/m2 4.6 µmoles photons m2/s
Table 7 Approximate equivalences between photometric, radiometric and quantum units
 44

Water and nutrients absorption by plants.

Water absorption by plants.

Almost all plant processes are related to water supply. The metabolic activity of cells and
plants overall is entirely dependent of water.

Water constitution

Is the water content that exists in plants. Over 90% of the living structures of the cell,
protoplasm, consists of water, the walls of all living cells are impregnated with water and
much of the volume of these cells is occupied by vacuoles filled with cell sap, water with
various dissolved substances.

Water constitutes 80% or more of the fresh weight of most herbaceous plant’s areas and
over 50% of the fresh weight of woody plants.

PLANT’S PARTS SPECIES WATER CONTAINED %

LEAVES Lettuce 94.8
Sunflower 81

STEMS Asparagus 88.3

Sunflower l 87.5
Pine-tree 45 – 70
FRUITS Tomato 94.1
Watermelon 92.1
Strawberry 89.1
Apple 84.0
SEEDS Sweet corn, edible 84.8
Dry corn 11.0
Table 8 Water content of various plant structures, % of fresh weight.

Coefficient of perspiration or relative consumption.

Growing plants transpire large amounts of water, and the amount of water in kilograms
consumed by a plant to produce one kilogram of dry matter, not including the roots, is
called relative consumption.

HELLRIGEL (TEMPERED) RISLER TEMPERED) YANKOVICH (SEMIÁRID)

CROP RC CROP RC CROP RC
 45

Wheat 399 Corn 216 Tomato 570

Oatmeal 402 Oatmeal 250 Corn 336
Pea plant 290 Zucchini 750
Pepper 810
Table 9 Relative Consumption for different regions (kg water/kg MS)

The RC as its name says it does not depend solely on cultivated species, but on the region
cultivated, growing season, soil type, moisture content and soil nutrient status.

SOIL FERTILITY WITHOUT FERTILIZER WITH FERTILIZER

POOR SOIL 550+- 16 350 +- 8
MEDIUM SOIL 479 +- 11 341 +- 4
FERTILE SOIL 392 +- 6 347 +- 6
Tabla 10 RC of corn cultivation at different levels of soil fertility.

The water constitution and the constant "loss" of water by plants through transpiration
must continually be compensated by means of absorption and transport from soil. The
"lost" can be considered a necessary evil since it has many beneficial aspects. Like the
nutrients supply dragged by the flow from soil nutrients, its distribution within the plant, as
well as the regulation of internal temperature of the plant.

Water enters the plant through the leaves, stems and other organs but most of it
penetrates the root, root hairs in particular represent a quite large exposed total surface2,
in addition to its higher permeability compared to older roots. The old roots do not have
hairs; they are suberized and are restricted to the minimum water inlet.

This is one of the main reasons for all cultivation from which a maximum performance is
intended to obtain, keep producing young and healthy roots throughout the life cycle,
either maintaining or improving soil conditions for their proper conduct or by stimulating
the plant in ordert to produce this type of root.

Due to the absorption process the plant replaces water loss by transpiration. Imbibition,
transpiration stress, metabolic absorption and osmosis: four distinguished types of force
that may explain the entry of water through the root.

2 It has been found that a single rye plant has about 14 billion root hairs with a total surface area of 370m 2 and the
rest of the root system of absorption totaled about 230 m 2. In contrast the leaf surface area was 5 m2 in
comparison to the stem area.
3 Suberin covers the cell walls and this makes them waterproof
 46

a) The entry of water imbibition into the cell is due because the protoplasm has great
affinity for water and their micelles adsorbes it easily; but once the protoplasm is
saturated, its ability to adsorb water finishes. It is not an important way of water
inlet into the root, although it may become so if the plant has been under
conditions of drought for a long time and is partially hydrated, and has high
importance in the hydration of seeds.
b) The perspiration tension force is present in the plant, caused by water evaporation
from leaves and steam removal through the stomata causes the generation of a vacuum
in the conducting vessels causing a water "current" towards stomata through the
xylem; consequently this force is an important but indirect absorption factor.
c) Active or metabolic absorption means that water is being absorbed with an
expenditure of metabolic energy, through a non-osmotic mechanism, this means it
moves against the diffusion gradient that is to say not an accelerated speed
(Kramer, 1959), for which it uses some of the energy from respiration, so that
absorption is only a minor fraction 5% of the total absorbed (Garcidueñas, 1978).
Finally this has not been proven conclusively.
d) Passive or by osmotic processes absorption does not require directly any energy
consumption, water penetrates from soil into the root in response to a potential
gradient which must continually decrease from the soil to the atmosphere. Plant
acts as a link in the water system in order to allow the downward flow of a potential
gradient from the soil to the atmosphere,

Water Potential,
The water in liquid state is a fluid whose molecules are in constant motion. The mobility of
these molecules depends on its free energy, ie the fraction of the total energy that can be
converted into work. The scale used to express and measure their free energy condition is
the water potential (Ψ). A water body, pure, free, without interactions with other bodies,
and normal pressure, corresponds to a Ψ = 0.

The Ψ is essentially determined by pressure and water activity. The latter depends in turn,
on osmotic effect, the presence of solutes, and the matrix effect, or interaction with
colloidal solid matrices. The Ψ can be expressed in terms of its components:

Ψ=Ψp+Ψπ+Ψm

The Ψp, pressure potential, is null at atmospheric pressures, positive at above atmospheric
pressures when hydrostatic pressures exists, and negative when there are tension or
vacuum conditions of atmospheric pressure. Also called gravitational.
 47

The Ψπ, osmotic potential represents the decreased water displacement capacity due to
presence of solutes. As the solute concentration (ie, the number of solute particles per unit
of solution volume) increases, Ψo becomes more negative. Without the presence of other
factors altering water potential, water molecules from solutions will move from places with
low solute concentration to areas with higher solute concentrations. The Ψo is considered
0 for pure water.

The Ψm, matric potential, represents the degree of water retention due to interactions with
solid or colloidal arrays, it can be worth 0 if there are no interactions or it can be negative.

Temperature affects the Ψ, an increase in temperature has a positive effect on its value and
a decrease tends to decrease it. The Ψ predicts how water will move under different
positions. Water moves spontaneously from a large area of water potential to an area with
the lowest potential, irrespective of the cause that causes this difference. Pressure is
another form of water potential.

The water within the cell

Water can be found in the cell wall and protoplast (mainly in the vacuole) of the plant.

The water inflow and outflow of protoplasm will depend on the relationship between his Ψ
and the Ψ of the external environment:

 If Ψinternal = Ψexternal: will be dynamic balance; no net flow.


 If Ψinternal > Ψexternal: there will be a net outflow of protoplasm, being able to reach
 the state of plasmolysis4.
 If Ψinternal < Ψexternal: there is a net inflow of water and consequently an increase in
volume of protoplasm, reaching the state of turgor

Water in the atmosphere.

The Ψ atmosphere is related to the relative humidity5 due to the water in the atmosphere
being vaporous with a very low pressure. Only HR values close to 100% condition the Ψ.
The Ψp in the atmosphere is 0.0 because P is atmospheric. Under these conditions the
water potential in the atmosphere at any temperature can be determined by measuring the
relative humidity.

4
Plasmolysis: process whereby plant cell loses water to be placed in a more concentrated she external environment.

The water in soil.

 48

The water in soil is subject to different force fields, so its potential differs from the one that
maintains pure water in its free state. Such forces result from the external gas-gravitational
attraction.

Thus, the full potential of ground water is the sum of the action of these factors
independently.

Ψtotal= Ψm+Ψπ+Ψp

 The gravitational potential (Ψp) is independent of soil properties, is determined by

the elevation of each point within with reference to an arbitrary level. Such level
and the point at issue within the soil are considered at the same height, in such way
the potential is always small or negligible, but if waterpower at soil surface is
compared with the force water has at 20m depths the Ψp would be large enough to
be taken into account.

 The matric potential (Ψm) plays an important role in water movement in soil. This
potential is a dynamic property closely related to forces of absorption and
adsorption for each soil, which vary with texture, bulk density, organic matter,
porosity and water quantity; thus, for example, a clay soil holds more water than a
 sandy soil at the same potential energy.
 This potential is also known as soil tension and can be measured with tensiometers.

5 Relative Humidity measures the relation between the quantity of water vapor (V) in a mass and the
maximum amount of water vapor this mass might support at a given temperature
 49

SANDY

CLAYEY

SILTY
LOAM
Figure 25 Characteristic retention water curves (g water/g soil)
for various soils, depending on matric potential Mpa

 Osmotic potential (Ψπ) results from the presence of solutes in the soil water, which
affect their thermodynamic properties, and decreases its potential energy. Is
important in saline soils where depending on the salinity, a possibility in which
water potential is lowered to such an extent might exist where there is no flow into
the roots and in extreme cases the flow direction is reversed. The total osmotic
potential is considered comprised by two components, one associated with free
solutes (soil solution) and the other that is retained (adsorbed) on the surface of
electrically charged colloids.

According to the force with which water is retained in a given time, it is classified into three
types:

a) Gravitational water, water that infiltrates by gravity to the deep layers,

Ψsoil=-0.0005 Mpa6
b) Pore water, water that remains retained by soil particles and remains available for
absorption by roots, and is exposed to evaporation.
Ψsoil=-0.03 Mpa
c) Permanent wilting point, amount of pore water that may no longer be absorbed by
roots. Ψsoil= -1.5 Mpa

Of course if soil conditions are suitable for growing a crop, the greatest potential influence
for water retention in soil is matric potential.
 50

6
1 bar equals 0.987 atm and at the same time 0.1 Mpa Water

transport in the plant.

Water movement in the plant, out of some exceptions is ascendant.

Theories explaining the ascent of water:

a) Radical pressure theory. When transpiration is significantly reduced or zero as

occurs during the night, and root cells can still secrete ions into the xylem. Since the
root vascular tissues are surrounded by endodermis, ions tend not to exit the xylem,
generating high solute concentration and decreasing the potential and water is
moved into the xylem by osmosis from the surrounding cells. This creates a positive
pressure that forces water and dissolved ions up through the xylem, in some cases
may cause accumulation of water on leaves, which is a notorious event at morning,
it is a fairly limited phenomenon and occurs mainly in short size plants.
b) Theory of capillarity and imbibition, as explained above.
c) Tense transpiration coheso theory. Refers to tensions created by the evaporation
and subsequent hydration of cell walls area (with the consequent movement of
water into the plant), is performed by a phenomenon of cohesion among water
molecules. When water evaporates anywhere in the plant (eg, leaf cells) the
reduction of water potential in the evaporating surface causes movement of the
latter from the xylem to the evaporative surface. This reduces the xylem water
potential, generating a negative pressure or tension. The negative pressure is
transmitted through the hydrodynamic system up to the root surface where the
water potential is decreased and causes the water influx from soil.
The origin of water evaporation in the stomatal cavities is the water potential under
dry air surrounding the leaves.
 51

Perspiration
Leaf
substomatal Location
spaces
Superior Air
Epidermis
Nerve
Laqunar
parenquima
Conductive element Leaves
Stomata

Conductive
elements
Perforation that
separates two
elements of the Vessel
same vessel
Bordered pits

xylem Stem

Conducting
vessels
Xylem root

Soil (root)
Water + mineral
Epidermis and
salts Soil
absorbent hair

Superior
Epidermis

Specialized
parenquima

Bundle
sheath
Tracheid
Sieve tube

Laqunat
parenquima
Apoplast
Inferior
epidermis
Air

Figure 26 (left) Water potential representation at different places on the path followed by water from the soil to the
atmosphere through the plant. (Right) Water potential from the xylem to the atmosphere through the leaf mesophyll.
 52

Due to the above reasons the water moves through the cortex, endodermis and pericycle,
until it penetrates the primary xylem, caused by potential differences. Two routes for the
water path in the root (Figure) are identified:

1. The symplast: set of protoplasts interconnected by plasmodesmata

2. The apoplast: set of cell walls and intercellular spaces.

In general it is considered that the apoplast composed mainly of cellulose and other
hydrophilic substances, exhibit a lower water resistance than the symplast wherein lipids,
hydrophobic substances, organelles and particles that increase viscosity abound. Water and
solutes can follow both routes, but it is believed that water flows via the apoplast pathway
wetting walls and intercellular spaces, reaching the endodermis where water and solutes
must continue via the symplast pathway and subsequently water follows via apoplast until
it reaches the xylem.

Epidermis The Caspary-2 Band

Protoplast
(SIMPLAST) Root exterior
Water
Cortex Cell walls
Endodermis
(APOPLAST) C aspray
Water Plasmodesmata Band
Endodermis Transverse
Root

Pericylce wall
Caspray Band interior
Reaching the endodermis, wáter and materials coming from
Xylem
apoplastic pathway (1) can’t keep going due to the Caspray
vesselsBand. They are required to penétrate into protoplast of
endodermic cells and continue through simplastic pathway
until crossing this layer (2). Water and materials coming via
Absorbent hair Caspray Band
Intercellular spaces simplast are not altering the pathway (3)
(APOPLAST)

The importance of this is knowing that water will have to go through the membranes and
its proper functioning requires ATP, any factor that negatively affects respiration at this
point, will affect the flow of water and nutrients.

Factors such as lack of oxygen caused by poor handling of water in the soil, structure
problems, salinity compaction, or mechanical or low soil temperatures will originate a low
or no respiratory rate and low energy production. On the other hand, the lack of water in
the soil or high salinity levels in soil would reduce the Ψsoil and limit the water entry into
the plant. The environmental temperature is another reason to limit water entry into the
plant to such an extent that limits transpiration by stomatal closure, the lack of light or low
transpiration by low temperatures.
 53

It is known that the solute influencing the opening and closing of stomata is potassium, as
its concentration increases the cells of stoma opens and when decreases the cell closes,
consequence of increasing or decreasing its volume by decreasing or increasing its Ψ when
water enters or leaves the cell. Chlorine is also involved and a substance that is synthesized
within the cell (malate) derived from starch hydrolysis, synthesis that will not be possible if
there is not enough phosphorus within the plant.

Light stimulates stomatal opening, CO2 is required to produce malate, but combined with
high concentrations of it at low luminosities causes stomatal closure.

Temperature greater than 35°C causes stomatal closure in most species, as respiration
increases and the intercellular concentration of CO2 increments.

Water content of soil and plants. If water loss by transpiration cannot be replaced by
absorption, guard cells lose turgor and the stomata closes before the leaf wilts. This
depends on the action of the "abscisic acid" hormone, which upon activation from root
cells causes the loss of guard cell K, with the consequent closure of stomata.

Stomatic cell Cellulose

microfibrils
Nucleous
Water

Ostiole

Chloroplast
Epidermic cells Fused walls Longitudinal
expansion

Figure 27 Stomata opening and closing.

Perspiration besides being the mechanism that results in water tension of the xylem and its
ascent in the plant also enables the distribution of water and mineral nutrients absorbed by
the roots throughout the plant.

Plant nutrient absorption.

Plants mineral absorption is possible due to its capacity to absorb inorganic ions at low
concentrations and to the extensive root surface. After the absorption are stored,
metabolized, or transported via xylem throughout the plant. The general shape of the root
system is defined by genetic mechanisms, but their proliferation is determined by the
availability of water and minerals in the surrounding environment. Mineral salts that are
more readily available are those found in a solution.
 54

The transport pathway comprises ion absorption by the roots and their movement through
the xylem to be transported to different areas of the plant cell:

a) Short-distance transport. Soil ions are absorbed by the root (passive and active). At the
root, they reach the endodermis where Caspary band interrupts the flow so they must
actively enter into the central cylinder where they will be transported. The transport
mechanism to the xylem usually is via apoplast or it can be directly absorbed by the
epidermis in root hairs and transported via symplast into the xylem.
b) Radial transport. Apoplast cylinder can be penetrated by ions and water from the
external medium. In a way that the plasmalemma of all cortical cells would act as a
large active surface for absorption into the root symplast.

Although there are exceptions, it is postulated that the final step of transferring the symplast
ions into the xylem is an active process, mediated by carriers, although regulated differently to
active absorption of the cells plasma membrane of the epidermis and cortex root.

Xylem
Cortex Trail

Active
absorption
Pathway
Apoplast
Caspray Bands

Figure 28 Model of apoplastic and symplastic pathways for radial ion transport through the root to the xylem.
Marshner (1986)

Nutrimental access mechanisms.

They are a series of processes and mechanisms through which nutrients move to the root
surface in order to be in the position of being absorbed.

1. Mass flow. The mass flow is a nutrients passive transport to the root along with
environmental water as it is absorbed and transpired by the plant. The force driving
mass flow, water plus ions toward the root is the water potential gradient
 55

established between soil water and water from the atmosphere through the plant.
Depending on the amount of transpired water from the solution concentration will
be the amounts of nutrients that make contact with the root surface.

FM=Va*C, water volume per element concentration (mol/m2/s)

2. Diffusion. Is a spontaneous process that leads to a net movement of a substance in

a high concentration region to another region where the substance. Diffusion is the
result of random thermal movement, of solute and solvent molecules in a solution,
or gas in the case of air. The net motion is most likely to move from an area of high
concentration, to the low concentration area. Thus the causing force of movement
is the concentration gradient.
D (mol/m-2/s)= Ion X diffusion coefficient concentration gradient.
DIFFUSION COEFFICIENTS OF SOME IONS IN SOIL
DIFFUSION (cm2/s)
ION
10-6—10-7
NO3-
1.4 x10-6
+
NH4
1 x10-8 – 2.4 x10-11
-
H2PO4
1.4 x10-6 – 9.5 x10-7
+
K
0.9 -4 x10-7
2+
Ca
0.9 -4 x10-7
2+
Mg
1.2 x10-6
-
Cl
0.5 – 8.4 x10-7
MoO42-

3. Radical interception - exchange per contact. As roots grow they have contact with large
soil volumes, pushing particles to the root sides, directly reaching nutrients that can be
absorbed. According to the surface area of the root volume, the concentration of
nutrients and the volume explored in the growth medium will be the contribution of
the IR-IC to nutrient absorption. On average soil conditions make contact between 1
and 2% of radical exploration, however in terms of container (pot), the IR-IC may be
the main mechanism of nutrient access, due to the large root mass per soil unit.
 56

When the supply of nutrients by mass flow and diffusion becomes limited, the plant
adopts a variety of strategies to optimize nutrient acquisition and is very likely that
under such conditions the IR-IC increases. Which is quite common for microelements.

NUTRITIONAL CORN SUPPLY DEVELOPED IN A FERTILE GROUND (Tisdale

NUTRIE NT
et al, 1990)
QU AN TITY

PERCENTAGES PROVIDED BY
ABSORBED MASSES DIFFUSION RADICAL

Table 11 Nutrients proportion in nutritional supply form

Relationship Between Nutrient Access And Fertilizer Location (Malavolta et

al, 1989)
CONTACT MECHANISM
Type of fertilizer
ELEMENT Interception Mass Flow Diffusion application

N 1 99 0 Distant
P 2 4 94 Localized
K 3 25 72 Localized
Ca 287 760 0 Broadcast
Mg 57 375 0 Broadcast
S 5 95 0 Distant
B 29 1000 0 Distant
Cu 70 20 10 Localized
Na 50 10 40 Localized
Mn 15 5 80 Localized
 57

Mo 10 200 0 Broadcast
Zn 20 20 60 Localized

Table 12 Nutrimental access and fertilizer position.

Mechanisms of nutrimental absorption. (Passive and active)

proteins

energy
Transported molecules

Figure 29 Types of Ion absorption and low molecular weight molecules.

A. Passive transport. It is a diffusion process of substances through the membrane.

Always occurs down the gradient, i.e. from a high concentration environment
towards the lower concentration. Can be given by:
1. Simple diffusion through a doble layer. In such way lipidic molecules enter as
hormones, steroids and non-polar substances such as oxygen and atmospheric
nitrogen. Some small size polar molecules, such as water, CO2, ethanol and
glycerin, may also pass through the membrane by simple diffusion. Water
diffusion is called osmosis.
 58

2. Simple diffusion through channels. Is performed by the so-called channel

proteins. Ions such as Na+, K+, Ca2+, Cl- enter through them. Channel proteins
are proteins with an internal channel hole, whose opening is regulated, for
example by another signal or a ligand that binds to a specific region in the
receptor channel protein which undergoes a structural transformation that
induces channel opening (figure below).

Transported
ligand
molecule

Channel rgulated by a ligand

Figure 30 Simple diffusion through channels

3. Facilitated diffusion. Allows the transport of small polar molecules such as

amino acids, monosaccharides, etc., that being unable to cross the lipid bilayer,
for their hydrophilicity, require transmembrane proteins to facilitate its passage.
These proteins are called proteins transporters or permeases, by binding to the
molecule to be transported undergo a change in structure that drags such
molecule inside the cell (flip-flop coupled proteins).
 59

Figure 31 flip-flop Coupled Proteins

B. Active transport. In this process transmembrane proteins act also, but these require
energy in form of ATP, to transport molecules across the membrane. Occurs when
the transport is performed against the electrochemical gradient.
There are three types of membrane proteins: catalytic proteins, proteins that form
solutes channels, and transport proteins. Catalytic proteins (enzymes) use energy to
pump protons (H+) through the membranes, enzymes that catalyze ATP are the
most abundant, Atepeasas, Proteins pumping ions against this gradient are called
ion pumps.
a. Transporters
b. Electrogenic or protons pumps.

A membrane capable of handling energy and attach it to do work is called energy-

transducing membranes and in plants there would be three: chloroplast membrane,
mitochondrial membrane and the plasma membrane or plasmalemma

a) Transporters Theory
 60

Figure 32 Transporters absorption model.

b) Electrogenic or protons Bombs

Concentration
gradient Concentration

gradient

Cytosol

Figure 33 Electrogenic or Protons Bombs

It is noteworthy that an important constituent of cell membranes is Ca ++ in order to

maintain its structural integrity. With calcium deficiency, they lose their ability to transport
solutes inside and are unable to retain them, that is, selective transport mechanisms are
disrupted, given their antagonism with some ions, besides membrane permeability
increases indiscriminately. The antagonistic effect mentioned also functions as a safety
mechanism for cells, for example against sodium.
 61

Overall, most species spend most of their biomass to aerial regions, suggesting that nutrient
absorption is controlled in part by processes occurring in the aerial region, in two senses:

 As demand the aerial region can increase the nutrients absorption in the root
making quick use of such salts in order to allocate them into growing areas

 As a contribution, the aerial region provides carbohydrates through phloem to
produce the ATP needed for nutrient absorption.

Factors affecting the absorption of ions.

 Physical and chemical characteristics of ions.


The constituents of membranes (phospholipids, sulfolipids and proteins) contain
electrically charged groups, and these groups interact with ions, the interaction force
increases according to the following sequence: uncharged molecules <monovalent
cations <monovalent anions <divalent cations <divalent anions <trivalent cations
<trivalent anions, while the nutrient absorption rate decreases in that order.
o Diffusion order in aqueous solutions
  
Cations: K+ > Na+ > Ca2+ > Mg2+
 
Anions: NO3- = Cl- > SO42- > H2PO4-
o Movement through the membrane
  
Cations: NH4+ > K+ > Mg2+ > Ca2+ > Na+

Anions: NO3- > Cl- 2-
> SO4 > H2PO
4-
 Cations > anions
 Interactions between ions.

Ions absorption requires their binding in specific transporters sites. It may have
 competition to some extent for occupying these places between ions with similar
 physicochemical properties (valence and ion diameter). It has been observed for
example that the NH4+ competes for binding sites with K+ affecting its absorption rate,
although the opposite does not occur. The affinity of Mg ++ (highly hydrated) for
 binding sites with transporters of the plasma membrane is particularly low, so that
other cations such as Ca ++, Mn ++, and even K + compete strongly with it
consequently its absorption rate can be greatly reduced.

Between NO3- and Cl- there is also competition. Greater external concentration of
Cl-less NO3- absorption rate and vice versa. This power can be used to reduce the
 excessive nitrate concentration in crop leaves of such as spinach, chard and lettuce,
 as consumption of nitrates and nitrites are considered harmful to health.
Furthermore, in saline substrates, the toxic effect of Cl- can reduce significantly
increasing the concentration of NO3-.
 62

Mg Absorption (µeq per 10 g of fresh weight per 8 horas)

Organ MgCl2 MgCl2+CaSO4 MgCl2+CaSO4+ KCl
Root 165 115 15
Aerial Region 88 25 6.5
Table 13 Potassium and Calcium Effect in Mg absorption in barley seedlings, concentration of each cation 0.25 me/l,
Marshner (1986)

Concentration (me L-1) Concentration in Leaves

- -
Cl NO3 (me Cl-/100g dry weight)
10 1.25 90
10 2.50 51
10 5.00 34
10 7.00 19
Table 14 Relation between the supply of nitrate and chlorine content in soybean leaves. Weigel et al. (1974)

Ions can also act synergistically. Stimulation on the uptake of cations and anions and
vice versa is often observed and is a result of the need to maintain a balance of
electrical charges within the cells. The clearest example of synergism is the stimulation
of Ca++ in both cation and anion absorption. This effect is due to maintaining the
stability and integrity of the membranes by Ca++, as this cation reacts with phosphate
groups of phospholipids keeping them together by certain force.

Nutrient excess Induced deficiency

N K
K N, Ca, Mg
Na K, Ca, Mg
Ca Mg
Mg K, Ca
Ca B
Fe Mn
Mn Fe
Figure 34 Common antagonisms
 63

  Cation: Anion Ratio

At low concentrations the absorption of a cation is not generally affected by its
 accompanying anion or vice versa; however, at high concentrations, the ions with
lower rates of absorption as SO4-- and Ca++ may negatively influence the absorption
of his companions ions.
Absorption Rate (µeq/g fresh weight/hour)
K+ of Cl- of
Concentration (me L-1) KCl K2SO4 KCl CaCl2
0.2 1.6 1.6 0.8 0.7
2.0 2.7 1.9 2.0 1.0
20.0 5.7 2.2 4.3 2.1
Table 15 Companion ion effect over the absorption rate of potassium and chloride in corn plants, Lügte and Latics
(1966)

The different absorption rates of cations and anions require a compensation of

electrical charges from both inside the cells and in the external solution. At high
external concentrations the charges of compensation within cells becomes the
limiting factor for ions absorption such as K + or Cl- when accompanied by SO4- and
Ca++ respectively. Cation excessive absorption may lead to an increase of pH in the
cytosol, however this pH must remain stable for proper enzyme functioning.
Therefore excess cations are compensated by an increase in the synthesis of organic
acids (R-COO-), with this subsequent transport of cations in the vacuole. Conversely
excessive absorption of anions tends to lower the pH of the cytosol due, among
other causes, the anion-proton symporter through the plasma membrane, causing
an increase in the decarboxylation of organic acids stored in the vacuoles.
A 70% of the cations and/or anions absorbed by plants are or represented by
ammonium or nitrate. The way in which the nitrogen is supplied has a different
effect on the organic acid content of plants and the solution pH to the outer
membranes. The increase in pH in the external solution for the nitrate absorption
results not only from the preferential absorption of anions but also of the nitrate
reduction produced by OH- according to the following equation:

NO3- + 8e- + 8H+ ----> NH3 + 2H2O + OH-

The OH- radicals can neutralize the H+ protons within the cytoplasm or may be
ejected into the external solution. In any case the pH of the external solution is
increased, which promotes better absorption of cations than anions in cytosol.

CATIONS ANIONS
Source of N Organic
Ca++ Mg++ K+ Na+ Total NO3- H2PO4- SO4-- Cl- acids Total
-
NO3 107 28 81 5 221 1 26 25 25 162 239
 64

NH4+ 72 22 49 7 141 1 25 25 31 54 136

Table 16 Influence in the form of nitrogen supply on the cation balance: anions in mustard. Kirkby (1968).

  External Concentration.
As already mentioned, the absorption of various ions is a predominantly active
process therefore at low concentrations follows a kinetic saturation (similar to that
of enzyme saturation by the substrate). If an ion concentration is increased the
absorption rate also increases but at a much lower rate (lower selectivity). The
 following figure shows the effect of the concentration of potassium and sodium on
their rates of absorption. Differences in absorption rate for K+ and Na+ reflect
differences in affinity of binding sites of the protein transporters in the plasma
membrane (high affinity for K+ and low affinity for Na+). The phosphorus absorption
curve is similar to potassium curve whereas calcium and magnesium are more like the
one of 8 sodium.
Absorption Rate (umol/g fresh weight/hour

6 Potassium

Sodium
2

0
1 2 3 4 5
Concentration (mM)
Figure 35 Potassium and sodium absorption by increasing its concentration in barley roots. Marshner, (1986).

  Internal concentration and nutritional status

The internal ion concentration is an indicator of the nutritional status of a plant and
an important regulator of absorption rates thereof. In general when the ion internal
concentration particularly increases, its absorption rate decreases and vice versa.
This is the result of several negative feedback mechanisms: ejection or efflux of ions,
changes in the number and activity of transporters, transformation rates and
incorporation of ion metabolism, charging speed of xylem and phloem, long
distances transport rates, vacuolar concentration, hormonal action, etc. These
regulatory mechanisms allow cells to maintain relatively constant nutrient
concentrations despite a wide range of outdoor concentrations.
 65

  Solution pH.
This determines the solubility of the ions or compounds formed in the soil solution.

The competition between H+ and other cations and between OH- and other anions is
very important for plant nutrition. A typical example of the influence of pH on the rate
of cations absorption is that there is a marked decrease in potassium absorption

with the increasing concentration of H+ (pH decrease between 7 and 4); this decrease is
caused by the competition for binding sites between H+ and K+ for binding in the
protein transporter of the plasma membrane. In contrast anion absorption is favored at
acid pH on the outside of the membrane. The effect of pH on the absorption of

nitrogen depends on whether this is supplied as NH4+ or NO3-. NH4+ is absorbed
faster than NO3- in alkaline pH and vice versa.

FORMATION OF COMPOUNDS OF CALCIUM AND PHOSPHORUS IN TERMS OF pH

NUTRITIONAL SOLUTION (Furlani, 1999)
pH nutritional solution
4.5 5.0 5.5 6.0 6.5 7.0 7.5 8.0
Forms Percentage of compound formed

Ca2+ 91.0 90.9 90.7 89.9 61.7 47.4 43.9 42.9

Ca SO4 7.2 7.2 7.1 7.1 5.3 4.3 4.0 3.9
Ca PO4 1.8 1.9 2.2 3.0 32.9 48.3 52.0 52.8

HPO4- 94.9 94.3 92.7 88.3 33.4 7.2 1.6 0.4

Ca PO4 5.0 5.3 6.2 8.5 63.2 91.2 97.8 99.2

Table 17 Effect of pH on the formation of certain compounds.

 66

 Plant selectivity. Each species has a preference for a particular type of ion in order
to fulfill the requirement of each phenological stage. We can appreciate it indirectly
by measuring changes in the pH of the soil solution or substrate.

Potassium

Nitrogen

Weeks after plantation

Figure 36 Ion Selectivity

  Salinity.
High concentrations of salts in solution as well as sodium and chloride directly influence
the absorption of nutrients. The Na directly affects the absorption of K and Cl affects
that of NO3. Osmotic pressure of the solution caused by the high salt concentration
decreases the gradient of water potentialof the plant, which results in a decreased
absorption of water. This is the reason why the absorption is affected by the nutrients
 that are transported by mass flow (perspiration) such as calcium and
 boron. To reduce or avoid these negative effects, electrical conductivity should be
between 1.5 to 3.0 dS m-1, which is equivalent to an osmotic pressure between 0.5-
1.0 atmospheres, where the plants can develop smoothly. Osmotic pressure is
calculated with the following relations:

Osmotic Potential (PO, kPa) = -42 CE -0.16 CE2

PO (atm) = -CE (0.36)
In both cases the CE in dS m-1

  Oxygen concentration
Root cells need energy from ATP for the absorption of the active ions. ATP is
synthesized in the respiratory process of such cells, process that requires oxygen.
When the oxygen concentration decreases a lot at cellular level, the absorption of
ions like potassium and phosphorus decreases markedly. Therefore a nutrient
deficiency is one of the major factors limiting the performance of growing crops in
soils with poor aeration.

 Temperature
 67

Respiration and consequent production of ATP is a temperature dependent process,

as well as an enzyme activity. Since enzymes are electrogenic pumps, it is clear that
within certain limits, at higher temperature there would be a higher rate of
absorption. At temperatures below 10 ° C, the root respiration rate is very low and
active absorption almost stops; Additionaly, in that temperature order, the plasma
membrane is less fluid featuring a greater resistance to the passage of solutes.

  Available carbohydrates and light

The main source of energy for cell respiration in the root are carbohydrates supplied
by leaves; if these are limited (eg for electricity restrictions in crops under high
population density), the rate of active absorption of ions is reduced.

Transport of nutrients via Xylem

Once the nutrients are "culled" or secreted into xylem, their movement in the same direction
begins (unidirectional) occurring by mass flow. During the day it comes in response to the force
of perspiration and is based on the forces of adhesion and cohesion of water (theory
explained). At night the transport is generated by the high concentration of solutes, root
pressure from respiration, this mechanism helps eliminate air bubbles that could have formed
during the day by cavitation. Xylem sap in addition to water, contains a large amount of
inorganic elements, some amino acids, organic acids and hormones, particularly cytokinins
produced in the root apex, also transported by this pathway abscisic acid. The pH is acidic.

Root pressure. The membrane permeability to water is greater than that of ions, therefore
the root cells behave as osmometers. The accumulation of ions in the apoplast of the wake
increases both the osmotic potential and the water potential, which induces a net flow of
water from the external solution. As a result of this flow the mentioned potential
decreases, while the hydrostatic pressure, or turgor pressure of the individual wake cells
increases. This hydrostatic pressure induces a flow of solution (water, ions and molecules)
in the xylem vessels into the stem; radical pressure is the given name of this pressure.
 68

Factors that affect ion transport in xylem.

a) External Ion concentration. External increased concentration results in increased ion
concentration in the sap.
SPEED OF ION MOVEMENT

ABSORPTION

DIFUSSION

EXTERNAL ION CONCENTRATION

Figure 37 external ion concentrations.

b) Temperature. In the long term due to changing growth patterns, such as a low
radical temperature will cause limited root growth, this creates limitations for intake
of water and nutrients. The low radical temperature is effective immediately in the
reduction of water permeability in roots, and in turn the water supply to the leaves.
A radical increase in temperature causes increase in the potassium concentration
but a decrease in xylem calcium concentration, which will cause changes in
membrane selectivity with temperature.
 69

Dry Weight (g plant)

Root Temperature (ºC)

K2
30
25 K1
Dry Weight
(g) of
the Plant

20
15 15 20 25
Sustrate Temperature (°C)
 70

Figure 38 (left) Effect of substrate temperature and potassium concentration in plnant's dry weight (K1=4me/l, K2=12
me/l), Chu and toop, 1975. (Right) Effect of roots temperature and P concentration on dry weight of the cultivated
tomato plant in sand. (P1 = 1 mgP/l, P2=5 mgP/l, P3=10 mgP/l).

c) Respiration. As the oxygen level in the soil decreases, the ion concentration in xylem
decreases too although the ratio between potassium and calcium remains.
d) Carbohydrate levels in the root. The raw material of root respiration is
carbohydrates accumulated in the root and the ions release into the xylem depends
on this energy. Photoperiod length affects the level of carbohydrates in the root.
e) Water. The effect of water deficit on the movement of water through xylem is
explained by the gradients that are created with changes in soil moisture.

f) Perspiration. The greater perspiration the greater movement of ions by the increase
in the mass flow of the external solution.
g) Adsorption and exchange. Although the mechanism of solute transport in the xylem
sap, by mass flow in not alive xylem vessels, occurs between the solutes, the cell
walls of the vessels and surrounding xylem parenchyma cells. The main interactions
are adsorption by polyvalent cations exchange and the release of organic
compounds by living cells. This cations adsorption and exchange is also present in
the cell walls of tissues.
h) Release or discharge. The composition of the sap can change along the path
followed by the release or secretion of solutes from the surrounding cells. The
secretion of nutrients from the xylem parenchyma is necessary to maintain a
continuous supply of nutrients to the growing parts in the aerial region. In periods
of ample supply of roots, nutrients are resorbed from the xylem sap and in nutrients
deficit periods they release them into the sap.
 71

Transport via phloem.

Phloem transport. Is bidirectional. Several theories have been proposed to explain the
phloem transport being one of the main the one related to mass flow (pressure flow theory
or turgor pressure gradient) based on the source-demand relation. Phloem sap is a lot more
concentrated than the xylem sap. The main components are carbohydrates, following are
the amino acids, organic acids, some mineral elements, especially potassium and boron
which are very important for the adequate phloem transport, some hormones such as
gibberellins are also transported, the auxin. Its pH is alkaline.

Factors affecting phloem transport.

a) Water. Reduction in the leaf water potential due to lack of water, induce a decrease
in the amount of similars entering the phloem, due to decreased photosynthesis
and photosynthate movement into the phloem.
b) Light, by affecting photosynthesis indirectly, besides influencing the movement of
sap inside the sieve tubes. Light can reduce starches and mobile carbon recycling on
leaves.
c) Temperature. Has a direct influence on the distribution of assimilates to affect either
supply and demand or the regions between them, according to their temperature.
d) Photosynthetic capacity. The position and leaf age determines the number and address
for the photosynthate. Young leaves mainly export to the apex of the stem, leaves and
branches. The older leaves export primarily down to the base of the stem.

e) Leaf position (phyllotaxis). The leaves located one upon another connects directly
with vascular rows, suggesting that export takes place following the most direct
vascular route.

Alternate Alternate Opposite Decussate

Whorled Grouped Fasciculate Interleaved

 72

SOIL.

The term derives from the Latin ground Solum which means ground or terrain and refers to
the "loose" surface on earth, to distinguish it from the solid rock.

"Soil is an unconsolidated mineral material on the immediate surface of the crust; A)

that has been subjected and/or influenced by genetic factors and from environment such
as the mother material, climate, macro and microorganisms and topography, all of them
acting for a while and causing a product --soil-- that differs from the material which is
derived in many properties and physical characteristics, chemical, biological and
morphological. B) From the point of view of plant growth "Soil is an unconsolidated mineral
material on the immediate surface of the crust that serves as a natural environment for the
growth of plants", able to provide support, water, organic and inorganic substances and air;
essential for crop development.
 73

Illustration 1 Diagram of the processes of pedogenesis

During the formation of earth’s crust there was a lot of volcanic activity and burial of plants and
animals. The volcanoes scattered lava and magma over earth’s crust transforming this into
various materials, and buried living creatures decomposed in various chemical elements.

The mineral part of the soil proceeds from various types of rocks.

1. Igneous rock: rocks formed by cooling and solidification of volcanic magma; is the
most abundant group of rocks on the Earth's crust.

2. Metamorphic rock: igneous rocks derived from sedimentary rocks and geological
disturbances.

3. Sedimentary rocks: Rocks derived from the destruction of other rocks exposed at
the surface. Sediments or product of such destruction accumulate in low areas
where they undergo consolidation or cementation.

Igneous rock

metamorphic
rock Heat and pressure
Siltstone

Disintegration, deposition and

foundation

Two processes of high importance in soil formation:

a. Weathering: The influences of the atmosphere and organisms on mineral rocks of the
Earth’s surface (dissolution, hydrolysis, carbonation, oxidation-reduction) Physical
effects of water are through percolation, capillary, erosion and physical chemical
flooding.
 74

 Physical Weathering: freezing and thawing, heating and cooling, wetting and
drying, erosion and action of plants and man.

a. Chemical weathering (decomposition):

a) HYDROLYSIS, water is an active substance H of the water tends to be

changed with the cation of various substances forming new
compounds, this occurs especially with basic compounds: CaSiO3
+ 2HOH → H2SiO3 + Ca(OH)2

b) HYDRATION: The binding of water with soil minerals, such as iron

oxide hydration. → 2Fe2O3 + 3HOH 2Fe2O33HOH the first named
compound is hematite (prints the ground red and the result is called
limonite (imparts a yellow color to the soil).

c) CARBONATION: Decomposition of organic matter in soil and plant

respiration releases large amounts of CO2 into the atmosphere from
the ground. This gas is readily combined with bases producing
carbonates and bicarbonates such as Ca (OH)2 and K(OH) derived
from hydrolysis: Ca(OH)2 + 2CO2 → Ca(HCO3)2 and K(OH) + CO2 →
KCO3 + H2O, this process is one of the most effective in decomposing
soil minerals. Since Ca and Mg bicarbonates are sparingly soluble.

d) OXIDATION: Oxygen freely combines with many other elements, so:

4FeO + O2 → 2Fe2O3 is the oxidation of ferrous iron in the ferric state
(Hematite), the S is easily oxidized to H2SO4

e) REDUCTION: This process is opposite to oxidation and occurs

whenever the O2, a common electron acceptor in the soil is found in
limited quantities. 4FeO + O2 ← 2Fe2O3
 75

f) SOLUTION: The water acts as a solvent in conjunction with CO2 or

with trace mineral and organic acids (HNO2, HNO3 , H2SO4 , H2SO4).

Continuous
Wheathering minerals Desintegration (size Primary resistant
slow (i.e. quartz, decrease) minerals (i.e. Quartz)
moscovite)
Desintegration in Minerals
Chemical and physical
(physical desintegration) Wheathering minerals alteration Silical clays
slow (i.e. teldespates
and biotite)
Decomposition and
Rocks (igneous, Wheathering minerals Other Silical clays
recombination
sedimentary, easy (i.e. calcite, sugite)
metamorphic)

Decomposition
Decomposition,

oxidation and Fe and Al oxides

hydtratation
Decomposition (chemical
desintegration)
Organic Complexes of

Solution
Organic acids solution
Al3+ and Fe3+
Solution

hydrolisis
Silical acid

Soluble Minerals
Solution

Illustration 2 Physical and chemical weathering

 76

Fussion

igneous rock
wheathering
metamorphism Metamorphic rock
Sedimentary rock

SOIL
Sedimentary rocks
FORMATION
formation
Erosion and
transport
Processes
Illustration 3 Diagram of soil
formation Sediments Sedimentation

Each of the stages and forms of weathering on a specific rock gives as result different
composition materials.

Name Composition
Primary minerals
Quartz Silicon dioxide
Feldspars Orthoclase Potassium aluminosilicate
Plagioclase Sodium and calcium aluminosilicate
Pyroxene Augite Fe, Ca and Mg aluminosilicates
Amphibole Hornblende Complex aluminosilicates mainly from Ca, Fe and Mg.
Micas Muscovite Hydrated potassium aluminosilicates
Biotite Iron and Magnesium hydrated aluminosilicate
Olivine Magnesium and iron silicate
Secondary minerals
Calcite Calcium carbonate
Dolomite Calcium and Magnesium Carbonate
Clay minerals Hydrated aluminosilicates - compounds
Limonite Hydrated iron oxide
Opal Hydrated silicon dioxide
Table 18 Primary and secondary minerals derived from weathering.

Each of these materials is rich in a mineral important in plant nutrition as shown in the
table below.

MINERALS
 77

PRIMARY SECONDARY

FOUND IN IGNEOUS AND METAMORPHIC FOUND IN SEDIMENTARY ROCKS FORMED

ROCKS AT HIGH TEMPERATURES AND HIGH AT NORMAL TEMPERATURE AND NORMAL
PRESSURE PRESSURE
QUARTZ Yes ALUMINOSILICATES (CLAY)
FELDSPARS K, Al, Na, Ca OXIDES AND HYDROXIDES
MICAS K, Al, OH, Fe, Mg CARBONATES
Fm SILICATES. Ca, Na, Fe, Mg, Al, SULFATES
Zn,
NO SILICATES Ca, P, OH, F, Cl. CHLORIDE
PHOSPHATE, ETC.
Table 19 minerales y compuestos derivados de las rocas.

2. Humification: The transformation of humic substances from primary organic

material. This process will be discussed in more depth later.

The soil layer is formed vertically, approximately parallel to the surface, called HORIZONS,
which differ in their morphological and physicochemical characteristics. The smaller volume
of soil whose lateral dimensions are sufficient to recognize horizons and study the
formation process called PEDON (pedion).

Finally by its physical and chemical soil characteristics it can have different uses, in such a
way that it may be consider that "The soil is the basis of all terrestrial ecosystems, therefore
life on earth depends on it."

Other soil uses:

1. As habitat

2. As a supply and water purification system

3. As recycling system

4. As building element

5. As a historical record
 78

From an agricultural perspective is important to know its current features, of

course, acquired by the process that originated but nowadays also by the handling it
has been subjected in order to achieve such result.

General composition of soil.

The soil comprises three phases:

1. Solid: Includes organic (living and nonliving) and inorganic components

2. Liquid: Water and soil solution. The soil water is not in a pure state, containing
substances dissolved or suspended and which together constitute "the soil
solution." Salts are dissolved substances in different ionic states and suspended
substances are from colloidal nature. The importance of soil solution lies in the
plants uptake of necessary nutrients and water from it, so that the composition will
directly influence the development of a plant; it is also important in the processes of
soil formation. The composition of the solution depends on several factors such as
lithology material that gave rise to the soil, climate (temperature and precipitation),
rocks solubility level, level of microbial activity in the soil, adding organic fertilizers
and mineralized amendments, and the quantity and quality of water used for
irrigation on agricultural soils.

3. Gas: soil atmosphere. There are essentially O2, CO2, N2, the renewal of these gases is
given mainly by gas exchange caused by the processes of respiration of soil
microorganisms and plants and gas exchange between the atmosphere and the soil.

The three phases must be interacting on such a balance that allows the plant to obtain
the resources it requires for its proper physiological performance and for ultimately
achieve higher performance or as aforementioned the maximum dry matter
production. This balance is based on each of the phases has the proper composition and
allow the action of the other phases, ie the solid phase should have a structure that
allows the proper flow of gases and air quality in the soil atmosphere is optimal for root
respiration and the proper growth and development of them, therefore of the entire
plant; and the quality of the liquid phase allows the soil to maintain this structure and is
a suitable source of water supply and nutrients to the plant; soil conditions in a given
time depends on the condition of each of the phases. Fortunately there is a component
of the solid phase that helps achieve a balance on these three phases because it has
multiple beneficial functions that directly and indirectly influence soil phases. This
component is organic matter.
 79

ORGANIC MATTER.
They are called organic compounds those whose basic element is carbon, and includes all
plant materials and dead animals as well as their transformation products found in and on
the soil. Two processes give decomposition of organic matter:

1. Mineralization. Transformation of organic matter by action of soil microorganisms in

soluble or gaseous inorganic compounds (SO42-, NO3-, CO22-), which can be used by
plants. CO2. Is mineralized carbon. Immobilization also occurs when these
compounds are used and integrated into the structure of soil microorganisms.
2. Humification. Compounds originated from the decomposition and are non-
mineralized become used to form new molecules of complex colloidal nature and
dark color, with high carbon content in their structure, which together constitute
the humus. The humidification process comprises the transformation of the
products produced from the breakdown of high and low molecular weight
molecules, from tissues of higher plants, animals and microbial cellular constituents,
as well as the molecular synthesis of these compounds.
Quality and Speed
of Response

Organic Matter Evaluation of the HUMICROP 100 effect on the Hass

Avocado
Gram sor cubic centimeters per plant

easily decomposable slowly decomposing materials

materials (eg manure) under anaerobic conditions

Mineralization Humification Absolute Fertilizer Fertilizer + Fertilizer +

Control HUMICROP 100 Control
Root Dry weight of
Volume the Plant
*Assessment made at commercial dose, witnesses were products
of similar origin and the incubation period was two weeks.
minerals without
formation of humus

Humus. The portion of the organic matter fully decomposed, and where there cannot be
differentiated any waste in particular. The compounds included under this heading are a
heterogeneous group of the following characteristics:

1. Highly polymerized substances (of aromatic and aliphatic compounds)

2. Relatively high molecular weight
 80

3. Dark color with colloidal and hydrophilic properties,

4. They are mostly composed of C, O, H, N, S and P.

Humus can be divided into:

1. Recognizable structures substances, amino acids, polysaccharides, fats and waxes.

(No humic substances)
2. Amorphous substances of complex structures, fulvic acids, humic acids and humin
himatomelanic acids. (Humic substances).

SOIL ORGANIC MATTER

VEGETABLE PROCESSED PRODUCTS:

WASTE HUMUS

RECOGNIZABLE AMORPHOUS
COMPOUNDS COMPOUNDS

POLYPEPTIDES FULVIC HUMIC

AMINOACIDS LIGNINS HUMINS
POLYSACCHARIDES ACIDS ACIDS

Because of its importance and influence on soil properties, we will describe the amorphous
compounds:

1. Fulvic acid: Portion of the soluble compounds of organic matter in water or dilute acids.
a) Molecular weight between 1000 and 5000
b) High content of carboxyl and phenolic groups
c) High interchangeability
d) Lateral and flexible structures
e) Ability to form stable compounds with polyvalent cations (iron, copper, aluminum,
etc.)
 81

CARBOXYL PHENOL
Figure 39 carboxyl group and phenolic groupfenólico

2. Humic acid is the soluble humus fraction in bases. This is because under alkaline
conditions the carboxyl groups are negatively charged by increasing the solubility of the
molecules, and precipitated in acid solution.
a) High level of polymerization
b) Molecular weight between 10000 and 100000
c) Low content of carboxyl groups (COO-), but higher content of aromatic groups (C6H5O-
)
d) Highly branched structure forming a spongy reticulum.
e) High water absorption capacity.
f) Less CIC than fulvic acids
g) Ability to form complexes
h) Because they are weakly dissociated they importantly influence in damping pH power
i) Very stable clay-humus complexes due to its ability to bind to clays.

Figure 40 Proposed structures of humic acids.

3. Humins: complex substances that are considered evolved from humic acids, which
union is so intense with the clay that has caused not dissolve in alkaline reagents.
a) High molecular weight
b) High content of oxygen relative to carbon
c) Low cation exchange capacity
 82

d) Aromatic condensed Groups

e) Structures resembling lignins

Humic and fulvic acids when applied to a floor can significantly increase the ability to
exchange cations, because the CIC proper to these compounds may represent from 2 to 30
times the CEC of clay loading constant. The cation exchange capacity of the organic matter
depends on the pH of the soil. Example: The cation exchange capacity of humus at pH 5 is
about 150 me/100 g, to pH of 6 will be 250 me/100 g.

CIC meq/100 g
500

40
20
5

HUMUS CLAY SILT SAND

Figure 41 cation exchange capacity of humus and different types of soil..

The organic matter content of soil usually expressed in soil weight percentage includes
organic matter on the floor in various decomposition stages based upon time and
conditions, either aerobic or anaerobic, which are subject to transitional organic, short-
term humus and stable humus originated under anaerobic conditions and that it can no
longer be decomposed by microorganisms representing approximately one third of total
organic matter.
 83

A soil containing more than 400 ppm of stable humus is considered fertile soil. To
accomplish such thing it is common to use materials that provide stable humus in small
quantities that generate large soil changes and often poorly understood by the extent to
which they are provided whilst the amount it represents relates to the total surface area,
but if the amount applied is compared to the area of influence on which the strong changes
are applied and can be noted in the concentration of stable humus in a small volume of soil
used for roots, in turn and observing significant improvements in soil.

SHORT TERM
ORGANIC MATTER HUMUS, 20%, 4
to 6 MONTHS

HUMUS DE
CORTO PLAZO,
20%, 4 A 6
MESES
STABLE
HUMUS,
35% 50 to MATERIA
250 YEARS ORGÁNICA
TRANSICIONAL,
TRANSITIONALO
45%, DESDE UNOS
RGANIC
MESES A VARIOS
MATTER, 45%
AÑOS
FROM FEW
MONTHS TO
SEVERAL YEARS

Figure 42 State of organic matter in soil (%)

 84

Process for humus extraction

NORMAL PROCESS
Leonardite

Alkaline extraction

Acid Extraction

HUMINS Humic Acids Fulvic Acids

Figure 43 Process for the stable humus extraction derived from leonardite.

Humus has wide influence on soil fertility, so that fertile soils are always associated with high
contents of humus in them, similarly it's productive capacity is increased with the addition of
humic materials due to its influence on the physical, chemical and biological soil (Table.)

SOIL PARAMETERS EFFECTS

Physical Favors the granular structure
Promotes ventilation
Facilitates drainage
Improves moisture retention
Increases the heat capacity
Reduces erosion
Chemical Regulates pH and salinity
Increases CIC
Enhances the solubility of nutrients
Source of N, S, and P
Reduces damage from pollutants
Biological Regulates microbial activity
Improves the establishment of the root
Promotes seed germination
 85

Improves crop nutrition

Table 20 Effects of organic matter on the soil

The effect of organic acids on the recovery of the structure of a ground is quite noticeable when
applying actually stable humus as to aggregation as soil particles into larger aggregates and
consequently the generation of larger pores (following figure), which together can improve the
ability of aeration, water movement in soil, water retention as well as a better development and
function of the cultivated plants root in order to have better environmental conditions.

UNALTERED SOIL RELATIVE CONTROL

TREATMENT STRUCTURE Porosity Aggregates
COEFFICIENT

UNALTERED
SOIL
RELATIVE
CONTROL

Figure 44 Humicrop 100 effect on the structure of soil Andosol. Dr. Victor Ordaz Chaparro, CP, Chapingo, Mex., CP,
Chapingo, Mex

The study unit is a soil profile or succession of layers called horizons; the way they are
arranged as well as the material affect both root growth (fundamental in plant nutrition
and healthy growth depends Body evolution of aerial parts), and water, air and nutrients
movement and retention, thus defining for each soil type specific physical and chemical
properties.
 86

PHYSICAL PROPERTIES OF SOIL

1) Depth. Thickness favorable for the penetration of ground roots of plants, it depends on
the volume of water that can be stored in the floor; is quite often the greater the depth
the greater density and lesser medium and large porosity. The root will deepen as far as
the conditions of aeration and drainage allow it to breathe properly.

Class Depth (cm) Description

1 0-30 Very shallow
2 30-60 Shallow
3 60-90 Moderate depth
4 90-120 Deep
5 >120 Very deep
Figure 45 Soil clasification based on depht

2) Texture. Relative proportion of sand, silt and clay in soil whose dimensions are smaller
than 0.002 mm clay, silt between 0.002 mm and 0063 mm and sand between 0.063mm
and 2mm. The importance of the study lies in the influence it has on the amount of
water that can be stored and retained in soil; water movement in soil; facility of water
nutrients and air supply.

a. Soil will have a high productive potential if the sum of fine silt and clay
(particles <10μ) represent a 40%
b. If the sum of fine silt plus clay is less than 40% the field capacity will be
reduced.
c. If the sum of fine silt plus clay is greater than 40% the aeration capacity will
be reduced.
d. The soil moisture is higher usable in silty loam soils than in sandy or heavy
clay soils
e. Sandy soils are less fertile than loamy and clay soils in terms of nutrient
content.
 87

Clayey

Clay-
Sandy- loam
clay
Franc-
Franc-clay-
clayey
Franc-clay-
sandy
Franc
Franc- Sandy Franc-loam
Sandy -
franc Loam
Figure 46 Triangle for Sandy texture
classification
WET AREA ON SOIL SURFACE
Sand percentage

Sandy Loam Clayey

CROSS SECTION OF WETTED ZONE IN THE SOIL

Sandy Loam Clayey

Figure 47 Wetting pattern and texture

It is considered that soil is:

a. Clayey if it contains more than 40% clay, small pores abound.

b. Loam if it contains more than 45% silt, balanced porosity.
c. Sandy if it contains more than 50% sand, abundant large pores.
 88

SAND SILT CLAY

RANK IN mm
SECONDARY (Colloid
DOMINANT PRIMARY
Mineral)
MINERALS
CHARGE LOW MODERATE HIGH
PROPERTIES

ATTRACTION BETWEEN LOW LOW- HIGH

PARTICLES MODERATE
ATTRACTION BETWEEN
LOW MODERATE HIGH
PARTICLES AND WATER

ABILITY TO RETAIN LOW MODERATE HIGH

IONS

HUMIDITY LOOSE SMOOTHE STICKY

CONSISTENCY MALLEABLE

DRY CONSISTENCY LOOSE POWDERY HARDD

Table 21 Relationship between texture, source material and other properties

Texture touch determination:

a. Clayey.- When wet, it adheres quite a lot, easily molded, the particles are not
visible and the surface shine slightly
b. Loam.- It sticks to the fingers, is molded with difficulty, fingers get greasy
appearance and the particles are brilliant.
c. Sandy.- Doesn’t sticks on fingers, is not shaped like a mass and individual
particles are visible.

3) Structure. Disposition or arrangement of fundamental particles of soil (sand, loam and

clay), forming larger units called aggregates that by joining together leave a series of
holes or helpful pores. The type of structure is determined by the overall shape of the
aggregates and the class is determined by the size of them: granular, laminar, and
subangular and prismatic.

The structure depends on the content of organic matter, calcium, sodium and magnesium
content but especially the relationship between these cations and wet conditions. Thus, the
granular structure is favored by high levels of organic matter, the laminar structure hinders root
penetration and promotes erosion, prismatic and angular structure denotes constant cycles of
contraction and expansion by drying and flooding respectively, finally the columnar
 89

structure is typical of sodic soils. Structure affects water penetration, drainage and aeration
thereby affecting root

development and productivity

Grain Blocky Plates

Rapid Moderate Slow

INFILTRATION SPEED
SOIL STRUCTURE
Granular Prismatic Soild

Figure 48 Structure and water infiltration speed

Structure formation:
 90

a. Flocculation: colloids tend to attract and stick together, flocculation occurs after
all the negative colloids are neutralized by adsorbed cations
b. Cementation or stabilization of flocules converts them into aggregates. Organic
polymers adhered or adsorbed on the clay surfaces function as glue or adhesive
avoiding their dispersion.
c. Humus organic polymers have 0.01-micron sizes and clay plates are one hundred
times larger (1 micron). This implies that many polymers are adsorbed onto the
clay surface and at the same time they interact with other polymers attached to
other clays.
d. These aggregates of clays with organic material to leave pores that allow
breathing soil and retain water.

Organic matter or humus that is interacting with the surface of the clay as part of the
aggregates is quite resistant to microbial degradation, since bacteria are often very large to
enter these spaces, besides being an anaerobic environment.

Other molecules that help particles keep their bound are oxides and carbonates, as well
as fungi and root exudates, also the roots of plants and fungi mycelia.
mucilage, Roots
flocculated carbonates,
colloids oxides, etc.

Silt, sand

Diagram showing two levels of organization of the particles in soil structure: the root is the one that
holds the larger aggregates at macroscopic level (soil ''lumps'’); lumps in turn are formed by
elementary particles joined by action of chemical bonds and organic exudates around the
flocculated colloids.

Figure 49
 91

The stability of the structure can be affected by:

a. Loosening. When dr aggregates are quickly wetted, an increase occurs in the inside
pressure and aggregates disintegrate.
b. Differential swelling rupture. When the clay swells when wet and shrinks when it
dries. This phenomenon is more severe in soils with higher clay content.
c. Mechanical disruption of aggregates. Mechanical rupture by destruction
mechanisms. Impact of raindrops, tillage, etc.
d. Physical chemical colloid dispersion. When the colloids tend to repel each other.
The principal agent of this is sodium, if the predominant ion exchange complex,
dominates by repulsive forces between colloids and structure it will tend to be
destroyed.

Regardless of degradation processes, stability of the aggregates varies under natural

conditions. The main soil properties that modifies them are texture, quantity and type of
clay, quantity and type of organic matter, quantity and type of cations that exist in the soil
solution and the presence of oxides and carbonates:

a. The higher clay contents the greater structure stability (up to about 50% clay).
b. Cations provide stability in the following order. Ca++> Mg++> Fe+++> Al+++ and
sodium on the contrary, favors the dispersion of the structure.
c. Some minerals give more stability than others: smectites> illite> kaolinite. And the
more polymerized the organic compounds are the more stability they provide.

4) Porosity. Represents the volume occupied by holes in the total volume. Consists of
different size pores, which act as follows:
a. Large pores: ø 0.01-0.05 mm. Ventilation and conduction line for the roots, supply
oxygen and evacuate CO2, through them the gravitational water is percolated and
aeration and infiltration improves.
b. Medium pores: ø 0.0002 to 0.01 mm. Store water and transport it by capillarity.
c. Small pores: ø < 0.0002 mm. Water not available to plants is stored.

The ratio of the sizes of pores is determined by the type of soil and the structure
compaction level, thus in clay soil will have a high amount of small pores while in a sandy
soil larger pores will predominate.
 92

The ideal soil must have 50% porosity, made up by 1/3 large pores and 2/3 medium pores.

The minimum soil porosity to allow good plant development is 35% in light soils and 45% in
heavy soils.

Estimates of porosity (EP%):

EP=[100(Dr-Da)]/Dr, where: Dr = Real soil density (g/cm3), Da = bulk density (g/cm3)

As well:

EP=(vol total pores/vol total soil) x100

Texture % Porosity
Thick sands 25 – 30
Very fine sands 30 – 35
Clayey sands 35 – 40
Silts 40 – 45
Clayey silt 45 – 50
Humic clays 50 – 65
Table 22 Porosity Ideal for every texture

5) Actual density. Refers to the average density of solid particles, independent of

structure. For a given horizon will remain more or less constant over time. An average
value for mineral horizons is 2.65 g/cm3 and 1.25 g/cm3 for organic horizons.
6) Bulk density. Refers to volumetric weight or weight per volume unit and comprising
solid particles in soil pores dry space. For same soil it can change relatively quickly,
depending on the compaction.

Material Da

Sands 1.6 – 1.7 gr/cm3

Francs 1.3 – 1.4 gr/cm3
Clays 1.0 – 1.2 gr/cm3
Organic Soils 0.7 – 1.0 gr/cm3
Table 23 Apparent density of different soil
 93

Texture Apparent Density g/cc % Porous space Soil weights

(Ton/ha)
0 - 25 cm 75 – 100 cm 0 – 25 cm 0 – 25 cm
Thick sands 1.84 1.86 26.3 4600
Clayey sands 1.56 1.65 37.51 3900
Silts 1.41 1.57 43.63 3535
Clayey silt 1.27 1.31 49.12 3175
Humic clays 0.92 0.98 62.93 2300
Table 24 Bulk density and porous space

Apparent density estimation:

Da=Pss/Vt, where: Pss= Weight of dry soil (gr), Vt= Total soil volume (cm3).

As well:

Da=[Solids Volume (cm3) X Real density (g/cm3)]/Total soil volume cm3)

Solids Volume = Total soil volume – Void volume (cm3)

The Da can be estimated in field by the following method:

Make a hole in the ground aproximetely of 20cm x 20 cm x 15 cm, the soil is removed from
said hole and is dried and weighed. The volume is calculated by placing a thin plastic over
the hole made and water is added to fill, quantifying the volume required for this.

Measuring the Da either in field or laboratory gives an idea of the soil compaction degree
and is useful for calculating water sheets, soil moisture and transformate ppm to kg/ha of
an element contained in the soil, by calculating the weight of the soil to a certain depth.

6) Hydraulic conductivity (cm/I). Soil ability to transmit water and oxygen indirectly in the
soil. This property is affected by the content of organic matter and exchangeable
sodium, these two together, modify the soil structure should apply that greater than 5
PSI invariably reduces the CH and therefore to improve soil substances that promote
improved soil structure by identifying the primary cause of degradation.

Northwest region of Mexico (Uvalle-Bueno, 1996)

Very Low Mild low Low Moderate High High Very High
moderate
 94

<0.04 0.04-0.42 0.43-1.67 1.68-4.2 4.21-12.6 12.61- >25.20

25.2

Bajio Region (Castellanos, 2000)

<1.0 1.0-2.0 2.1-3.0 3.1-5.0 5.1-10.0 10.1-20.0 >20

Table 25 Hydraulic Conductivity (cm/he).

In the following case, degradation of soil structure caused by the loss of organic matter
over time has affected substantially the hydraulic conductivity.

YEAR M.O. (%) N (kg/ha) P2O5 (kg/ha) C.H. (cm/hr)

1945 2.2 40 0 4.2

1955 1.9 80 0 3.5
1965 1.6 100 46 2.6
1975 1.4 120 46 1.4
1985 1.3 180 46 <0.4
1995 1.2 240 46 <0.4
Table 26 Soil behavior through time, CIANO 1996, Valle del Yaqui, Son.

7. Soil air. Air occupying free of water pores. The air outlet of soil O2 by the plant is a
condition for taking water and nutrients.

8. Soil respiration. Gas exchange occurs by diffusion caused by the difference in partial
pressure of O2 and CO2, between the atmosphere and the rhizosphere; this movement
is affected by temperature, atmospheric pressure, wind and precipitation.

O2 % CO2 % N2 %

Atmosphere 20.95 0.03 79

 95

Rhizosfere 20.6 >0.2 79

R. Compact Soil 10 10
Table 27 Concentration of gases in different environments

So the root can take nutrients, it must expend energy and energy will be obtained from
respiration.

• Content greater than 5% of CO2 and/or less than 10% of O2 affects root respiration
and plant growth. (Directly related to structure).

• CO2 in rhizosphere come 1/3 of root respiration, 2/3 of microorganism’s respiration.

• Moreover, CO2 excess generates higher concentration of CaCO3 and raises the
concentration of bicarbonate in the soil, causing nutritional or physiological
disorders.

9. Aeration capacity. Fraction of porous space occupied by air in the soil:

Aeration capacity (%) = Porous space (%) - Volume of water (%)

10. Drainage capacity. Ability to move in soil water at field capacity and its bases in texture
and structure.
Drainage capability (%) = Porous space (%) – Water volume in CC (%)
Va=Ps*Da

When the relationship between Aeration capacity/Drainage capacity is less than 0.5 would
result in nutritional disorders and when is greater than 2 to physiological disorders
preferably.

CHEMICAL PROPERTIES OF THE SOIL

a. Ground reaction or hydrogen potential (pH). Expresses the concentration of dissolved
H+ ions in the soil solution in relation to the concentration of hydroxyl ions OH- solution
dissolved therein. If the amount of H+ is greater than OH- then the solution will be
acidic and the pH value will be low, conversely decreases if the number of H + ions and if
OH-ions increases the solution will be alkaline.
The ratio between these two ions is given in a logarithmic scale.
1

= []

That is, that the rate of change between the ions H+ and OH- is 10 times its concentration for
each point of pH change, and soil pH 5 is 10 times more acidic than the same soil with pH 6.
 96

The pH value is between 0 and 14 and in agricultural soils we can find values between 4 and
10, and is considered neutral when its value is 7 and would be acids with less than this and
alkaline pH value greater than 7.

Ions that generates acidity are H3O+ y Al3+, hydrogen directly and Aluminum does by
passing the soil solution from the decomposition of water molecules:

i. Al3+ + H20 → Al(OH)2+ + H30+

ii. Al(OH)2+ + H20 → Al(OH)2+ + H30+,

iii. Al(OH)2+ + H20 → Al(OH)3 + H30+

At each step a proton is released causing a decrease in pH.

The Al3+ complex can bind to change complex by blocking gear positions and achieves a
reduction of the CIC. In soils where high amounts of basic cations exist this process is low,
because these cations are those that bind to water molecules.

i. Colloid-Ca2+ + 2H20 → colloid-H2 + Ca(0H)2

In calcareous soils it can cause acidity with microbial activity in wet periods and
temperature suitable for their work.

ii. H20 + CO2 → H+ + OH- + CO2 → H+ + HCO3

The H+ cations originating from this reaction causes a decrease in pH.

Furthermore, the induced decrease in acidity may be performed as follows

Soil Soil
(lime) (wáter (Carbon
dioxide)

Figure 50 Liming reaction.

 97

Liming reactions function as follows: The Ca2+ of lime is replaced by Al3+ o H+ in exchange
sites, and the carbonate ion (CO32-) reacts with soil water solution by creating an excess
ions 0H, which in turn react with the excess of H+ (acid), forming water.

Soils acidified as a result of removal of Ca++, Mg++, K+ y Na+ through crops leaching and
consumption. When cations are removed from soil they arereplaced by ions generators of
acid as H+ o Al+++. Nitrogen fertilizers also contribute to soil acidification, reacting in soil
increase the concentration of H+ in the soil solution.

The pH is considered an important indicator of fertility because the favorable effects of

reaction nearly neutral ground, this favors nitrification, symbiotic nitrogen fixation, and
good availability of most nutrients positively influence the crop growth. Considered optimal
pH range for most crops between 6 and 7.5, and if considered appropriate organic soils a
range between 5 and 5.5. Furthermore, the pH gives an immediate idea of the exchange
complex saturation.

Acidity Alkalinity
Extreme Very Strong Weak Very Very Weak Weak Strong Very
strong Weak strong

NITROGEN

PHOSPHORUS

POTASSIUM

SULFUR

CALCIUM

MAGNESIUM

IRON

MANGANESE

BORON

COPPER AND ZINC

Figure 51 Nutrients availability according to pH

 98

Excess acidity produce calcium deficiencies to plants and microorganisms, excess aluminum
or manganese in the soil solution generating toxicities for its high concentration and poor
or no root development, also has adverse effects on the humification and mineralization of
organic matter and fixation of atmospheric nitrogen.

In basic pH soils may also be problems of toxicity, microelements deficiency and possible
loss of soil structure.

Affects physical properties by inhibit grouping of particles to form aggregates, and chemicals, to
inhibit or reduce the ability to bind nutrients and biological to reduce the microorganism’s
activity and consequently modify transformation cycles of organic compounds.

Another concept that is handled respecting acidity is the potential acidity, and refers to the
total exchange positions occupied by H+, in equilibrium with a given concentration of H+ in
soil solution, therefore, depends on the soil colloids and CIC.

A tendency to compensate soil pH changes is known as a power buffer. Soils with high
damping power (High content of neutralizing cations in the exchange complex or high
carbonate content) will be able to maintain a roughly constant pH, with changes that occur
in the solution, ie, the greater the CIC is, greater the probability that there will be enough
change cations to neutralize pH fluctuations. Acid soils are well buffered against the bases
and basic soils are well buffered against acid.

When the soil is high in carbonates, neutralization occurs at the expense of carbonates
dissolution, generating calcium and soluble bicarbonates than will precipitate as calcium
carbonate when the soil dries.

iii. CaCO3 + H+ Ca++ + HCO3

b. Cation exchange capacity.

Is the measure of the ability of a soil or soil materials to retain exchangeable cations.
Can be defined as the amount of negative charges per unit of dry mass of soil that is
neutralized by exchangeable cations, or indirectly, the number of moles of positively
charged adsorbed ions, which can be exchanged per unit of dry mass, under some given
conditions of temperature and pH.
The unit of measure is in cmol/kg, or me/100g.
CIC is constituted by soil colloids, clay in its different types and organic matter in different
levels of decomposition. To the negative charges existing on the colloids surface can be
 99

fix cations from the soil solution. The positions to which a cation can be fix are called
shift positions. The adsorbed cations can pass from the solid phase to the soluble phase
and viceversa, according to several factors or rules.

CIC Rules

a. The positions of change tend to be always busy

b. The location of each ion tends to be balanced so that there is the same amount in
the exchange complex as in the solution.
c. Once in equilibrium, by increasing a cation concentration an amount of this tends to
fix on the exchange complex moving another that was already set. This implies that
some ions are fixed harder than others due to:

i. Its valency, if are two equal concentration cations its adsorbed the one with
higher valency; if two ions of equal valency in different but low
concentration, is mostly adsorbed the cation with lesser relative
concentration. This is because approaching the ideal concentrations, the
more diluted is the more activity has and will be joined electrically faster.
(Soil purifying action, Pb, Cd, etc.)
ii. Concentration, the higher the concentration of cation in the solution, the
greater the amount adsorbed, (regulates intoxication process).
iii. Hydration degree of the cation in soil solution. The cation is adsorbed that is
less hydrated. The hydrated has a larger ionic radius.
Cation adsorption order:
Al+++>>Ca++>Mg++>>NH4+>K +>H+>Na+>Li+
d. When an ion is displaced from the complex exchange for another that is attracted
more strongly, this remains in solution unti it can move to another that is fixed with
less force than that.

There are two considerations for the abovementioned:

a. Certain colloids negatively charges do not allow an easy ions exchange set therein,
for example the loads interlaminar clays.
b. Most loads on the colloids surface are negative, but there is also anion exchange
capacity, in which anions adhere.
 100

The exchange between solution and omplex change is always EQUIVALENT TO EQUIVALENT: If enter
two charges, two loads must leave, for example, by adding potassium to a calcium saturated soil the
following reaction occurs due to exchange processes, K+ may come to replace the Ca++.

i. SOIL Ca + 2KCl Ca2Cl + SOIL 2K

The arrangement of ions in the soil solution, relative to colloids, is explained by the theory of
the diffuse double layer. Around each colloid there is a double layer of charged ions: at the
nearest layer ions are retained more strongly, whereas with increasing distance to the
colloid attractive forces on the ions of the solution are weaker and concentration is smaller.
Therefore, the concentration of cations is strongly influenced by the behavior of the
exchange Against Positive Ion complex. Against Negative Ion

colloid highly
negative

Stern Layer

Diffuse Layer

Balance Ions with

Solution

Figure 52 Diffuse doble layer diagram

As mentioned in the previous topic, the pH gives us an idea of the kind of basic predominant
cation, acid (H+ o Al++) or basic (Ca++, Mg++, K+, Na+), the latter condition the existence of a
basic pH to promote H+ fixation to the exchange complex as noted in the following reaction.

i. Soil Ca++ + H2O Soil H2 + Ca(OH)2

As a result two concepts related to CIC arise:

a. Exhangeable bases Sum (S). Is the sum of me/100g of Ca++, Mg++, K+, and Na+ that
are adsorbed to soil exchange complex. The difference between CIC and S
represents the acidity change, is the sum of the acidity cation generators that are
linked to exchange complex (mainly H+ and Al+++).
b. Degree of base saturation (V). Indicates in what percentage the exchange complex is
occupied by exchangeable bases. V= S/CIC*100
 101

Figure 53 Level of base saturation and pH

The cation exchange capacity of clays depends on two factors: the surface charge density
and specific surface area.

Charge density is the amount of negative charges per unit area. This depends on the
amount of isomorphic substitution of elements, a certain positive charge, by other
elements less positive charge, in its crystal structure, remaining free in each case negatively
charge and charges are not compensated by the edge of the crystals.

Clay types and its chemical properties

a. Clays of constant charge occurs by isomorphous replacement.

i. Montmorillonite (2:1) the isomorphic substitution in the octahedral layer
where Al3+ is replaced by Mg2+
ii. Vermiculite substitution is in the tetrahedron, the Si4+ is replaced by Al3+

The charge of each mineral depends on the having amount of isomorphic substitution, and the
strength of the negative charge will depend on whether the substitution was made in the
octahedron (interior) or tetrahedron (exterior clay).
 102

Octahedron

Tetrahedron

Figure 54 Clay structure and isomorphic substitution.

a. Variable charge clays.- are iron oxides (goethite, hematite), oxides of Al (gibbsite),
and allophane, (amorphous aluminosilicate).
i. In these clays there’s no isomorphic substitution, the charge is carried at the
edges.
ii. Its charge is pH dependent, in acid environment the edges will be
protonated and wil develop a positive charge in alkaline environment will
lose these protons and develop a negative charge.
iii. Tend to form covalent bonds with P, immobilizing it on soil.
iv. Allophane is mainly found in volcanic soils and has no crystalline1 tubular or
spherical structure forming a large contact surface. It has the ability to retain
a lot of water

Negative charge Positive charge

Coloid Total Constant Variable (cmol/Kg)
Organic 200 10 90 0
Esmectite 100 95 5 0
Kaolinite 8 5 95 2
Gibsite 4 0 100 4
Table 28 Charge characteristics of some colloids.

1
Crystal structure means that the ions that conform it are placed in an order and this order is repeated in all
mineral.
 103

Organic
Colloid

Mineral Colloid pH depending

charges

blocked charges by
Permanent Al (oH)3
Charges

Influence of pH on CIC of an organic and a mineral colloid.

Figure 55 pH effect on CIC mineral and organic colloids

Negative Charge
Superficial Charge

Net Charge
Charge point Zero

Positive Charge

Soil pH
Figure 56 Volcanic soil charge

Specific surface. Is determined by the colloid surface in which cations can be fixed. As the
particles are smaller increases the surface area increases.

i. 1 g of Illite has a contact surface of 110 m2

ii. 1 g of Montmorillonite has a contact surface of 700 m2
iii. 1 g of Allophane has a contact surface of 1000 m2
 104

Secondary Union Swelling Specific

Mineral Type condition/ potential Surface

No interlaminar
kaolinite surface; strong Barely
(non-expandable) union nothing

Week unión,
Montmorillnite great high
(expandable) expansion
Week unión,
Vermiculite great high
expansion
(expandable)
Parcial K loss,
strong union
Hydrated mica low
(non-expandable)

Table 29 Specific surfaces of some clays

Thus, the colloids with high charge density and specific high surface area are those with
higher cation exchange capacity.

Organic matter as noted in the chart above and specified in the following, is subject to a CIC
change in function of pH to lose H+ as you increase the value of this.

Figure 57 pH dependent charge

Capacity significance of cation exchange. CIC knowledge is required for predicting the soil
behavior of organic or colloidal soil materials alone or added to the soil.

a. Is an important fertility rate, because it controls the availability of major cations for
plant growth: Calcium, Magnesium, Potassium, Sodium, and cations that regulate
other soil chemical property: Aluminum, Hydrogen, Ammonia.
 105

b. Plays an important role in the clays flocculation and dispersion process and thus in
the development of the aggregates structure and stability.
c. Determines the soil role as natural debugger to allow contaminant retention
incorporated into the soil, such as Pb, Cd, Hg.
d. Allows inference about clays mineralogy in each horizon.

Determining the need for soil cation.

There are three considerations involved in determining the need for each of the three
principal soil cations.

a. Cation exchange capacity. The different types of soils hold adsorption in different
total amounts of the cations sum, which depends on the type and amount of clay,
silt and sand and organic matter content.

Material CIC (cmolc/kg) Texture CIC

CLAYS Sandy 2-3
Kaolinite 3-15
Sandy Loam 3-10
Illite and Chlorite 10-40
Halloysite 5-50 Clear 11-16

Smectite (Montmorillonite) 80-150 Clear loam 13-26

Vermiculite 100-150 Clayey and slilty clay 4-58
Alophane (volcanic soils) <100 Table 30 CIC of different textural classes
Humic substances 300-500
Organic Matter 150-500
Oxides and hydroxides Fe and 4
Other materials
Silt quartz <0.01
Quartz 1-2
Feldspars 1-2
Table 31 CIC Clays and other materials

b. Ratio of such elements. CIC considers a fixed amount for a given soil, but not so
the ratio of adsorbed cations, which can be modified to achieve an appropriate mix
for the growth of certain plant or crop.
 106

Level BASE SATURATION %

VERY LOW Ca Mg K
LOW <50 <5 <1
MEDIUM 50-60 5-9 1-3
HIGH 60-75 9-15 3-7
Table 32 Base VERY HIGH 75-85 15-20 9-10 saturation levels,
% CIC MUY ALTO >85 >20 >10

CROP Ca Mg K TOTAL
(%) (%) (%) (%)

Tomato 70-75 8-12 3-5 81-92

Melon 75-80 8-10 3-5 86-95

Papaya 70-75 10-12 5-7 85-94

Avocado 65-70 9-12 6-8 80-90

Apple 65-70 12-15 4-6 81-91

Table 33 Base saturation by crop.

Cations Appropriate relationship

Ca/Mg 4– 8

Ca/K 11 – 25

Ca/Na > 18

Mg/K 2– 5

(Ca+Mg)/K 12 – 30
Table 34 Appropriate relationships between cations

C. Desired saturation degree. In most cases is not necessary and is not economically
profitable saturate the exchange complex with bases. Between 80% and 90% of CIC
saturation with a balanced proportion of exchangeable bases would be suitable for
most crops

Base saturation calculus.

With the cation exchange value and concentration of each of the bases, reported by the
laboratory, we can calculate the percentage of each in the exchange complex and decide
 107

according to the percentages suggested whether to supplementing soil to a cation or not.

The applied amount and the number of fractions into which must be applied is decided
according to the target saturation suitable for the crop; the chemical form of the compound
to be applied will depend on other soil parameters as pH, chlorine concentration and soil
sulfates and water to be used for irrigation.

The CIC is expressed in cmolc/kg or me/100g, and to calculate the bases saturation, they
should always be expressed under these units. It is common that the concentration of said
cations in the soil are expressed, in the analysis result, in parts per million, at which should
be transformed to me/100g by the following equation.
me cation ppm
100g from cation = equivalent cation weight x 10

An equivalent weight is equal to the atomic weight divided by the valency.

ELEMENT ATOMIC WEIGHT VALENCY EQUIVALENT WEIGHT

Ca 40.08 2 20.04

Mg 24.31 2 12.16

K 39.1 1 39.1

Na 22.9 1 22.9

Table 35 Equivalent bases weight

Then, percentage of each of the bases will be calculated with the following equation.
% = 100gme cation
CIC

In this way it is possible to compare the current bases saturation percent against suggested.

LIME TYPE FORMULA PNT

INCREASE PH
Calcium carbonate CaCO3 100
Dolomitic lime CaMg(CO3)2 109
Quicklime CaO 179
Hydrated lime Ca(OH)2 135
KEEP OR DECREASE PH
 108

Gypsum CaSO42H2O
Table 36 Materials that provide Ca and Ca+Mg, and how modify the pH

FERTILIZER FORMULA REACTION SALINE SOLUBILITY

INDEX (g/L)
Magnesium sulfate MgSO4 7H2O Neutral 44 710
Potassium chloride KCl Acid 116.3 700
Potassium sulphate K2SO4 Neutral 46.1 110 (30°C)
Potassium sulphate and K2SO4.2MgSO4 Neutral
Mangesium
Table 37 Fertilizers used for magnesium and potassium intake to achieve the balance of the soil bases.

c. Electrical conductivity of soil. It is an indirect measure of the concentration of dissolved

salts in the soil solution. This gives us an idea of the amount of dissolved salts but not
the type of salt dissolved, however, knowledge of this parameter shall quickly and easily
determine whether a salts concentration is beneficial or harmful for the crop in
question, solely on the basis of the amount of salt, since some crops are more tolerant
than others to high levels of salinity and dissolved salts in the soil solution.

The determination of electrical conductivity is important to make decisions on:

i. Land use for agricultural purposes.

ii. Soil handling
iii. Establishing a crop
iv. Irrigation sheets
v. Washing requirements
vi. Water handling in irrigation systems

Salts at high levels reduce the osmotic potential of the soil solution by reducing the
availability of water for the plants, even when the soil exhibits a proper humidity level.
Salts derived from irrigation water, and fertilizer added either directly to the soil or
dissolved in irrigation water; excessive accumulation of salts in soil comes from
mismanagement of irrigation and excessive application of fertilizers, or even if its a
suitable fertilizer and water application may exist any barrier that block the water
vertical flow and keep the salts in the depth where the roots are more active. It is
common to confuse the terms salinity and sodium. A soil can be saline, sodic or saline
sodic. Saline soils are handled with washing, sodic and saline sodic soils with application
of calcium-based breeders and subsequently washed.
 109

The electrical conductivity can be measured in saturation extract or in a soil: water ratio
1:2, at a temperature of 25°C, which is more practical and simple, however it is
desirable to know each other equivalents.

CE soil:water 1:2 Ces

<0.15 <0.4
0.15-0.5 0.4-1.2
0.5-1.0 1.2-2.4
1.0-1.5 2.4-3.8
1.5-2.0 3.8-5.5
2.0-2.5 5.5-7.9
>2.5 >7.9
Table 38 Relationship between CE in saturation extract and EC in the ratio 1:2, for soils of different textures in central
Mexico.

CEs SALINITY CONDITIONS PLANTS EFFECT

<1 Free salts soil There is no restriction for any crop

1-2 Soil very low in salts. Some very sensitive crops can restrict their yield.

2-4 Moderately saline soil Crop yields can be affected in performance

4-8 Saline Soil Performance of almost all crops is affected by this

salinity condition.
8-16 High saline soil Only very resistant crops can grow in saline soils.

>16 Extremely saline soil Virtually any conventional crop can economically grow
in this soil.
Tabla 39 Soil classification and its overall effect on crop from Ces..

Sodium particular case.

As already mentioned, although sodium is an essential element for plants, as found in high
levels in soil, causes various problems to both the soil and the plant, its effect can be
understood by expressing the content based on percentage that occupies exchange bases
saturation, therefore must be expressed as an exchangeable sodium percentage, PSI.
According to USDA, a soil is considered sodic from a sodium content of 15 sodium
exchangeable percent (PSI), but in practice it is observed that much lower levels in fine-
textured soils causing problems of physical order. In these soils, with higher levels of 6-10
PSI the structure starts to deteriorate and the sodium negative effect is not by specific
toxicity caused directly by the ion to the plant, but by the problems of water and gases
 110

impermeability without excluded that some crops dramatically reduces development at

these levels for PSI.

% BASES SATURATION
LEVELS Ca Mg K Na
VERY LOW <50 <5 <1 <1
LOW 50-60 5-9 1-3 1-3
MEDIUM 60-75 9-15 3-7 3.1-5
HIGH 75-85 15-20 9-10 5.1-20
VERY HIGH >85 >20 >10 >20

Table 40 Base and PSI saturation percentage

CLASSIFICATION PSI CROPS GROWING RESPONSE

Extremely 2-10 Deciduous fruit, walnut, Toxicity sodium symptom to low
sensitive citrus, avocado PSI
Sensitive 10-20 Beans Limited development to low PSI
Moderately 20-30 Clover, oats, fescue, rice Sodium response but with
sensitive and dalis grass. favorable soil structure.
Tolerant 30-40 Wheat, cotton, alfalfa, Limited development due to
barley, tomatoes. nutrition and structure factors

Very Tolerant >40 Beets, grass, Rhodes. Development usually limited due
to unfavorable structure.

Table 41 Tolerance of some crops to sodium level (as specific ion) on the basis of change, PSI.

When the sodium level is high sodium bicarbonate and sodium carbonate are formed,
producing a pH increase and may reach up to 10 under stringent conditions and generate
Fe, Zn and Mn deficiencies. A soil with pH greater than 8.2 in water, is usually a sodic soil
and requires rehabilitation. Overall in clay soils when the sodium level (extracted with
ammonium acetate) exceeds 400 ppm surely will have some sodium problems.

In a soil where there are several limiting productivity factors but there are high levels of
sodium, this problem need to be solved first. Rehabilitation is done by adding a source of
calcium and the most common is the agricultural gypsum (CaSO4.2H2O) or by adding
calcium "maker" such as sulfuric acid, ammonium polysulfide or calcium or sulfur. These
products react with soil calcium carbonate, produce calcium sulfate and works just like the
agricultural gypsum, with the only limitation for use, the soil must be lime, ie, having at
least 2% carbonate calcium; otherwise high addition of acid will collapse soil pH with dire
consequences for fertility.
 111

To calculate the amount of improver, PSI, CIC, and Da depth, data are required. To fine or
medium textured soils the goal is to return soil to a PSI less than 5%, although it will be
convenient to consider the crops presented tolerance to the sodium level in soil, especially
for those extremely sensitive.

The calculation is performed as the foregoing calculation of base saturation, whereas a

calcium milliequivalent (1 gypsum me) will replace a sodium milequivalente of the
exchange complex. A gyspumI me weighs 87mg, once the me to displace are calculated, the
amount of gypsum2 used is obtained. Doses vary from 1 ton to 50 ton per hectare
depending on the CIC and the problem severity; rehabilitation is carried out in several
months after enhancer application, when it is carried out the reaction, a situation that
requires an adequate level of moisture, and is advisable to perform a washlways after this
process, to remove excess sodium sulfate of soil.

CIC Exchangeable sodium content in soil, cmolc/Kg or me/100g of soil.

1 2 3 4 5 6 7 8 9 10 15 20 25
50 0 0 1.5 4.6 7.7 10.8 13.9 17.0 20.1 23.2 39 54 70
45 0 0 2.3 5.4 8.5 11.6 14.7 17.8 21.0 24 39 55 70
40 0 0 3.1 6.2 9.3 12.4 15.5 18.6 21.7 24.8 40 56 71
35 0 0.8 3.9 7 10.1 13.2 16.2 19.3 22.5 25.5 41 57 72
30 0 1.5 4.6 7.7 10.8 13.9 17.0 20.1 23.2 26.3 42 57 73
25 0 2.3 5.4 8.5 11.6 14.7 17.8 20.9 24 27.1 42 58 74
20 0 3.1 6.2 9.3 12.4 15.5 18.6 21.7 24.8 27.9 43 59 74
Table 42 Agricultural gypsum requirements applied to the soil according to PSI, to reduce it to 5%, considering Da = 1.2
ton/m3 and Pr = 30 cm.

2
One ton of pure gypsum equals to 186 kg of sulfur, 570 kg of sulfuric acid, or 660 kg of calcium
polysulfide.
 112

SALTS CEs TYPO EFFECTS RECOVERY

(dS/m)
LOCATION
y PSI
Nonexistent CEs<2,
PSI<7
Dissolved in solution CEs>2, Saline Salts affect mainly plants Salt Leaching
PSI<7 (difficulty to adsorb
water and toxicity) and
not soil properties.

Sodium adsorbed on CEs<2, Sodium Deteriorate soil structure Improved

the change positions PSI>7 chemistry,
calcium intake,
Sodium adsorbed on CEs>2, Sodium Dissolved salts avoid the Improve
the change positions PSI>7 saline destruction of structure chemistry and
and dissolved in problems. Similar effects salt leaching.
solution to those of saline soils

Table 44 Classification and management of soils based on their content of salts and sodium.
 113

Water in soil.
Given the need for water consumption by plants is important to know the relationship with
the environment where it is stored and where the plant uses it through the root, soil, and
promote the right conditions under which the plant make the least effort to obtain it.

As seen before, the spaces between soil aggregates form a network of interconnected
cavities in different shapes and dimensions. By adding water to a dry ground, this is
distributed by the cavities around the particles and it is retained by adhesive and cohesive
forces; displaces the cavities air, and finally, fills the pores.

Of the number of pores that have been filled and the force with which water is held we can
find different types of soil water.

a. Gravitational water. Water draining by gravity, this occurs when the pores are filled
and the film around the particles is of a thickness such that gravity force is greater
than the cohesive forces holding the film. This type of water generates undesirable
effects on soil depending on the texture, such as flooding, nutrient leaching and
decrease soil aeration. The drainage is greater in sandy soils than in clay soils.

b. Pore water. Once free water is filtered, water is stored in medium and small pores
and is still moving by capillarity. This is water available or usable by plants and is
between field capacity and permanent wilting point of plants.

c. Hygroscopic water. Having completed the movement of water by capillary action,

water is pinched tightly in the form of very thin layers, loses its liquid state and
moves as vapor, so that water is not usable by plants.
 114

Saturation
Gravity wáter

Rapid Drainage

Field capacity

Capilar wáter
Usable water
Slow drainage

Permanent
wilting
Hygroscopic wáter
Unavailable inexistent drainage
water

Figure 58 Types of water, its availability and drainage characteristics.

Soil moisture constants.

a. Saturation Point: Maximum water amount that soil can hold. The type of clay,
texture and organic matter content determines this point.
Usually determined in laboratory.
Texture %
Sandy Less than 10
Clayey and organic Until 150
Table 45 Moisture content at saturation

b. Field capacity: The water amount of that a soil can hold against gravity after irrigation
or rain that has soaked the entire soil. And it is estimated that water is retained in soil
at a tension of 1/3 atmosphere.

CC = (Ps saturation/1.84) -0.48

Texture %
Sandy 5-16
Loamy 15-30
Clayey Loam 25-35
Clayey 30-70
 115

Table 46 Moisture content at crop capacity, Palacios, 1966.

c. Permanent Wilting Point: Is the moisture soil content percentage in which the plants
wilt and can no longer recover. Water held at an equivalent tension to the osmotic
pressure in the plants roots, approximately 15 atmospheres however can oscilate
between 10 and 20 atmospheres, which is not important because this large tension
change occurs for very small change soil moisture.

PMP=CCX0.595.

The moisture constants are for calculate the maximum irrigation sheets to apply when
related to the apparent density and soil depth or roots, and to calculate the irrigation
frequency based on a fraction of the available moisture (HA) ie a fraction of the difference
between CC and PMP, which for pressurized irrigation and intensive crops, such fraction
shall be less than or equal to 20% of the HA.

HA=CC-PMP

TEXTURE USABLE MOISTURE

cm water/30 cm Soil depth

Very coarse sand 1.0-2.0

Sand and Sandy loam 2.0-4.0

Loam and slit loam 4.0-5.5

Clay loam, slity clay loam, 4.5-6.2

clay sandy loam.

Arcilla arenosa, arcilla limosa y arcilla 4.0-6.2

Table 47 Available moisture in different textures.

L= (%)− (%) X Da X h,
100
 116

Where: L= Irrigation sheet (cm) CC= Field capacity (%), PMP= permanent wilting point (%),
Da=Apparent density (a-dimensional3), h=soil or radicular depth

This is indeed the maximum amount of water that can be harnessed to a particular depth
or said in another way the maximum amount of usable water that can be applied to a soil
considering a certain depth and that irrigation will be giving when the soil is PMP. Since it is
not convenient to have a crop and afford reach PMP, only a fraction of the available
moisture is used and this must be replace according to the crop consumption that will
depend on the species, phenological stage and of course weather conditions

Between a dry and wet soil there is a very considerable variation in moisture content,
which is generally expressed in percentage by weight of dry soil.

That is:

Ps=(Psh-Pss)/Pss*100, where: Ps= Moisture content (% de dry soil), Psh=moisturized soil

weight (g), Pss= Dry soil (g)

Or for estimate the moisture content in volume percentage:

Pv=Ps*Da, where: Pv=Percent by volume (%), Ps=Moisture content in % on weight, Da=

Apparent density.

The moisture content in the soil may be obtained by the gravimetric method as stated
above or estimated in terms of the tension with which water is retained in the soil, through
the use of tensiometers or gypsum blocks and generally this is expressed in centibars 4 (cb),
however, this implies the need to know the curve of moisture soil tension and taking as
input the tension it is possible to know the moisture content of the soil.

Plant growth response to the moisture content in the soil is given as follows:

a. Deficient. Content less than PMP. Wilt dehydration, loss of cell turgor and plasma
destruction dryness. No growth after PMP.
b. Between PM and PMP content (6-15 atm), growth is reduced due to a low water
content, thereby breathing and also maduration accelerates. The root radical
system is relatively large.

3
Refers to the aparent density of soil, g/cm3, with respect to the density of water, g/cm3, which is a-
dimensional. This relationship is called apparent relative density or specific gravity and quantitatively is equal
to the aparent density of the soil.
4
1 atmosphere = 1.013 Bar
 117

c. Optimal. Profitable moisture (0.15 to 6 atm). Between these moisture levels the
optimum growth occurs and best quality, and this represents between 20 and 100%
of the available moisture in a floor.
d. High. High moisture contents are given a large vegetative development but little
root development, delayed maturation by high turgor that stimulates the formation
of new cells and there susceptibility to lodging.
e. Excess. Damage from lack of oxygen, excess CO2 and toxics. No water or nutrient
intake.

Water quality
The quality of irrigation water determines the suitable way to use, or the management of
soil moisture in relation to the physical and chemical properties of soils, crop tolerance to
salts, irrigation method employed, conditions of internal and surface soil drainage.

Water quality for agricultural crops irrigation, can be determined according to three criteria:

CRITERIA INDEXES SYMBOLS

Soluble salt content Electrical conductivity CE
Effective potential SE
Salinity potential SP
Probable effect of Sodium on Sodium adsorption ratio RAS and adjusted
the physical characteristics RAS
of soil. Residual sodium carbonate CSR
Possible sodium percent PSP
Content of toxic elements in Boron content B
plants. Chlorine content Cl
Sodium content Na
Table 48 Criteria and indexes classification of water irrigation. Palacios and Aceves (1970)

1. Soluble salt content. The effect of accumulated salts in soil can be harmful if the
amount becomes excessive so as to reduce the solution osmotic pressure probable
 118

surrounding plant roots, causing the plant to make greater efforts to take water and
after certain limit this can reduce performance or even die. High salt water requires to
be handled with an adequate irrigation sheet including in this certain fraction to
drainage and thus maintain salt balance in the soil. To realize the danger of soil salinity
from the use of a particular type of water with an x salt concentration at application,
the soil become saturated soil, salt concentration in this solution will be 5X and to reach
capacity field the concentration will be 10X, only by effect of salt concentration thereof
in less water. Estimating salinity parameters are described below.

1. Electrical conductivity. As in soil solution electrical the conductivity is an

indirect measure of dissolved salts in water. As increasing dissolved solids in
the water increases the ease of conducting electrical current and this is the
physical way to measure this variable.
The measure units are dS/m and are equivalent to mmhos/cm.
The CE can also be expressed in terms of total dissolved solids or TDS, mg/L.
1 dS/m is equivalent to approximately 640 mg/L, in the range from 0.1 to 5
dS/m. However, this equivalence may vary with the type of salt from 0.5g to
1.2 g/L per dS/m.
The sum of cations or anion, given in me/l, dissolved in water is equal to 10
time the CE in dS/m, ie 1 dS/m = Σ anions(me/l)/10 or Σcations(me/l)/10, in
the range of 0.1 dS/m to 5 dS/m.
Also a dS/m is equivalent to an osmotic potential of 0.36 atm, in the range of
3 to 30 dS/m, and this is what ultimately leads to the loss of plants, because
they spend more energy for water soil as CE increases.

CLASSIFICATION CEa to 25°C, dS/m

Low salinity 0-0.25
Moderate salinity 0.25-0.75
Medium salinity 0.75-2.25
High salinity 2.25-4.0
Very high salinity 4.0-6.0
Extremely high salinity >6.0
Table 49 Water classiffication for salinity, Richards et al (1954)

Based on this classification many laboratories indicate that when water has a salinity higher
than 2.2 dS/m should not be used in agriculture, therefore, this classification is questioned,
 119

since water is used up to 5 dS/m in an appropriate way, this is possible only if the waters
are calcic and no sodium.

MAXIMUM CE DUE TO VARIOUS SALTS

Maximum solubility
Salt in

Table 50 CE Maximum CE due to different salts.

The solubility of the salts depends on the solvent temperature in this case, water, so the
electrical conductivity measurement is standardized at 25 ° C
 120

Table 51 Maximum solubility in water of some salts found in soils at different temperatures.

Figure 59 Salts solubility according to temperature.

 121

SALT SOLUBILITY g/L

CaCO3 0.0131
MgCO3 0.42
Na2CO3 213.0
K2CO3 829.0
CaSO4 2.04
MgSO4 262.0
Na2SO4 185.0
K2SO4 108.0
Tabla 52 Salts solubility at 20ºC

2. Effective salinity (SE). It is a more realistic estimate of the danger presented

by irrigation water soluble salts to become part of the soil solution,
considering the subsequent precipitation of the less soluble salts, such as
calcium carbonate, magnesium carbonate and calcium sulfate, and thus
become ineffective in the osmotic pressure of the soil solution. This process
is more relevant when the water has a high content of carbonates and
bicarbonates.
For its calculation there are four possibilities:
a) If, Ca>(CO3 + HCO3 + SO4), therefore:
SE= Σcations- (CO3 + HCO3 + SO4)
b) If, Ca<(CO3 + HCO3 + SO4); but Ca>(CO3 + HCO3), therefore:
SE= Σcations- Ca
c) If, Ca<(CO3 + HCO3); but (Ca + Mg)>(CO3 + HCO3), therefore:
SE= Σcations- (CO3 + HCO3)
d) If, (Ca+Mg)<(CO3 + HCO3); therefore:
SE= Σcations- (Ca + Mg)
All values are expressed in me/l
3. Potential salinity (SP). When the soil profitable moisture decreases below
50%, the chlorides and some sulfates are the last remaining salts in solution.
Potential salinity is an index to estimate the risk of such salts and therefore
considerably increase the osmotic pressure.
Index is calculated using the following formula:
a) SP=Cl + ½ SO4
All ions are expressed in mg/l
 122

Clasification SP SE

Good <3 <3

Regular 3-15 3-15

Bad >15 >15

Table 53 classification, based on their salinity potential and effective salinity.

POTENTIAL SALINITY
SOIL CONDITIONS Clase 1 Clase 2 Clase 3
Poorly drained soils <3 3-5 >5
Soils with slow drainage <5 5-10 >10
Well-drained soils <7 7-15 >15
Well-drained soils, moderately tolerant crops <10 20-30 >30
Well-drained soils, highly tolerant crops <15 25-35 >35
Table 54 Water classification, based on their salinity potential and soil specifications.

a) Sodium probable effect on the physical soil properties. When the sodium
soil concentration is high and calcium and magnesium precipitate in the
soil solution by the action of carbonates and bicarbonates, then sodium
accumulates and replaces the calcium and magnesium in the exchange
complex, turning into an electrical imbalance of colloidal particles of soil
and due to the predominance of negative charges the particles repels,
aggregates deflocculate and the soil loses it structure, creating a
waterproof layer on the ground surface and in turn reducing the input
water and gas exchange, also promotes compaction, all affecting the
normal development of crops.
a. Sodium adsorption ratio. It is one of the most used indexes to
measure the danger of sodification soil, representing the irrigation
water. Is a relationship between the concentration of Ca and Mg,
and correlates with the exchangeable sodium percentage will have
the soil once equilibrate with the irrigation water. De acuerdo con
esto, entre mayor sea el valor de la RAS mayor será el PSI del suelo y
el peligro de sodificación. Accordingly, the higher the RAS value the
greater will be the soil PSI and the sodification danger.
[Na]

b) =
√
[Ca]+[Mg]

2
 123
100∗(−0.0126+0.01475∗RAS)
c) =

Where Na, Ca, and Mg are Sodium,

Calcium and Magnesium in meq/L from
wáter analysis.

Figure 60 Nomogram for determining RAS values of irrigation water and for estimating the corresponging values of PSI
from soil which is in balance with water. Bulletin 29 FAO, (adapted from Richards 1954).).

[Na]

d)= [Cax]+[Mg]
(Suarez, 1981)
√

The RASaj consider the total salt content and the bicarbonates presence,
which if not removed from the water precipitate calcium and this
contributes to the sodium problem is further aggravated, in addition that the
CO2 partial pressure in the soil surface.

Substituting the value of Ca by Cax in RASaj equation, which is obtained from a

table that relates the ratio HCO3/Ca and salinity in irrigation water in dS/m
 124

do the calculation. The RASaj, is a more accurate and recommended way to

predict the danger of sodium in soils by using water with high sodium and
bicarbonates contents.

Water salinity applied in soil, (EC)

(dS/m)
0.1 0.2 0.3 0.5 0.7 1.0 1.5 2.0 3.0 4.0 6.0 8.0
.05 13.20 13.61 13.92 14.40 14.79 15.26 15.91 16.43 17.28 17.97 19.07 19.94
.10 8.31 8.57 8.77 9.07 9.31 9.62 10.02 10.35 10.89 11.32 12.01 12.56
.15 6.34 6.54 6.69 6.92 7.11 7.34 7.65 7.90 8.31 8.64 9.17 9.58
.20 5.24 5.40 5.52 5.71 5.87 6.06 6.31 6.52 6.86 7.13 7.57 7.91
.25 4.51 4.65 4.76 4.92 5.06 5.22 5.44 5.62 5.91 6.15 6.52 6.82
.30 4.00 4.12 4.21 4.36 4.48 4.62 4.82 4.98 5.24 5.44 5.77 6.04
.35 3.61 3.72 3.80 3.94 4.04 4.17 4.35 4.49 4.72 4.91 5.21 5.45
.40 3.30 3.40 3.48 3.60 3.70 3.82 3.98 4.11 4.32 4.49 4.77 4.98
.45 3.05 3.14 3.22 3.33 3.42 3.53 3.68 3.80 4.00 4.15 4.41 4.61
.50 2.84 2.93 3.00 3.10 3.19 3.29 3.43 3.54 3.72 3.87 4.11 4.30
.75 2.17 2.24 2.29 2.37 2.43 2.51 2.62 2.70 2.84 2.95 3.14 3.28
1.00 1.79 1.85 1.89 1.96 2.01 2.09 2.16 2.23 2.35 2.44 2.59 2.71
1.25 1.54 1.59 1.63 1.68 1.73 1.78 1.86 1.92 2.02 2.10 2.23 2.33
Relation
HCO3/Ca 1.50 1.37 1.41 1.44 1.49 1.53 1.58 1.65 1.70 1.79 1.86 1.97 2.07
1.75 1.23 1.27 1.30 1.35 1.38 1.43 1.49 1.54 1.62 1.68 1.78 1.86
2.00 1.13 1.16 1.19 1.23 1.26 1.31 1.36 1.40 1.48 1.54 1.63 1.70
2.25 1.04 1.08 1.10 1.14 1.17 1.21 1.26 1.30 1.37 1.42 1.51 1.58
2.50 0.97 1.00 1.02 1.06 1.09 1.12 1.17 1.21 1.27 1.32 1.40 1.47
3.00 0.85 0.89 0.91 0.94 0.96 1.00 1.04 1.07 1.13 1.17 1.24 1.30
3.50 0.78 0.80 0.82 0.85 0.87 0.90 0.94 0.97 1.02 1.06 1.12 1.17
4.00 0.71 0.73 0.75 0.78 0.80 0.82 0.86 0.88 0.93 0.97 1.03 1.07
4.50 0.66 0.68 0.69 0.72 0.74 0.76 0.79 0.82 0.86 0.90 0.95 0.99
5.00 0.61 0.63 0.65 0.67 0.69 0.71 0.74 0.76 0.80 0.83 0.88 0.93
7.00 0.49 0.50 0.52 0.53 0.55 0.57 0.59 0.61 0.64 0.67 0.71 0.74
10.00 0.39 0.40 0.41 0.42 0.43 0.45 0.47 0.48 0.51 0.53 0.56 0.58
20.00 0.24 0.25 0.26 0.26 0.27 0.28 0.29 0.30 0.32 0.33 0.35 0.37
30.00 0.18 0.19 0.20 0.20 0.21 0.21 0.22 0.23 0.24 0.25 0.27 0.28
Tabla 55 Cax values to calculate the RASaj

1 Adapted from Suarez (1981).

2Asumes that the calcium source is CaCO3 or silicates; no precipitation of magnesium, and CO2 partial pressure on the soil
surface (PCO2) is 0.0007 atmosphere.

3 Ca , HCO3, Ca are reported in me/l; EC is in dS/m.

x
 125

Water with high RAS represents more risk to the ground than while the EC is lower. This is
because among the lower its higher electrical conductivity the higher will be the impact of
sodium in reducing the hydraulic conductivity and infiltration rates. However, take note
that on the other hand, salt water also possesses the risk of soil salinization and a negative
effect on crop yields. As already mentioned, the way to solve a salinity problem is by
washing and sodicity problem is by adding calcium-based breeders and calcium makers,
either the soil or water.

Severe reduction in the

infiltration rate Mild to moderate
reduction in the
infiltration rate
Adsorption ratio of Sodium RAS

Without reduction in the infiltration rate

Water applied salinity (CE) ds/m

Figure 61 Relative index of reduced water infiltration into the soil affected by salinity and RAS. Bulletin 29, FAO.
 126

Decreasing Permeability

Stable Permeability

Concentration
Ratio between critique concentration and interchangeable soil sodium percentage in
the center of Tezcoco river.

As a way to prevent the soil infiltration problems, caused by the irrigation water, should
maintain a minimum salinity in relation to RAS as follows:

D ) C= 0.35*RAS + 1.43
Where C is the minimum concentration or critique that must have
irrigation or washing water to prevent deflocculation, (me/l), always
verifying the following equation.
E ) C - CEwater, if the difference is negative the water is adequate and
there is no need for an amendment, if it is positive, then, to prevent soil
deflocculation the salinity must be changed.
2. Residual Sodium Carbonate, (CSR). This expressed as me/l, is another danger indicator
of soil sodicity and takes into account the calcium and magnesium precipitation by
carbonates and bicarbonates, leaving the rest of suitable carbonates or bicarbonates to
form highly soluble sodium compounds; therefore also decreases
the antagonistic effect on sodium.
a) CSR= (CO3 + HCO3)-(Ca + Mg)
 127

Clasification CSR

Good <1.25

Regular 1.25-2.5

Bad >2.5
Table 56 Water classification according to CSR

If there is a bad water respect to CSR, the way to solve this problem is by
using acids to destroy carbonates and bicarbonates and be replaced by the
accompanying anion, PO4, SO4, NO3, using phosphoric, sulfuric or nitric acid
respectively

TEMPERATURE °C CO3Na2 CO3HNa

0 70 68

10 122 80

20 213 93

30 371 107

40 441 121

50 429 137

Table 57 Solubility of sodium compounds at different temperatures (g/l)

pH of the saturated CaCO3 solublity in

solution with CaCO3 me/l
6.21 19.3
6.5 14.4
7.15 7.1
7.85 2.7
8.6 1.1
9.2 0.8
10.12 0.36
Table 58 Solubility at different pH
 128

3. Possible Sodium Percent (PSP). The danger of calcium and magnesium

displacement by sodium in the exchange complex begins when the sodium
content in the solution is more than half of the dissolved cations. Under these
circumstances the sodium proportion in solution is not sufficiently
representative of this danger as the less soluble salts precipitate and thus
increases the Na % relatively, so the PSP is more representative when referring
to effective salinity. All values are expressed in me/l.
a) = NaSE ∗ 100

It is considered that if the water has a PSP value greater 50% can convert a
soil to sodium soil through its application.

SOIL CONDITION PSP CLASSIFICATION

Any >50% Good for irrigation

Organic soils or light texture <50% or >50% Good for irrigation

but with less than
10 me/l of Na

Mineral soils of medium to >50%, with more Sodification hazard, water should be
heavy textures with less than than 10 me/l of mixed with others of better quality or
4% of CaCO3 + MgCO3 Sodium plaster must be added to them.

Table 59 Classification of irrigation water according to PSP

 129

Riverside Rules for the evaluation of irrigation wáter quality

Very high
High
Inde
Medium
Sodium

x
Low

Conductivity

Low Medium High Very high Excesive

Salinity
Good water quality suitable for irrigation.
Water quality suitable for irrigation with
precaution Water not for irrigation.

Figure 62 Diagram for classifying irrigation water for their CEx106 and its RAS.

Type Meaning of quality and use

C1 Low salinity water, suitable for irrigation in all cases. There may be problems
only in very low permeability soils.
 130

C2 Average salinity water, suitable for irrigation. In certain cases it might be

necessary to use excess volumes and salt tolerant crops.
C3 High salinity water, suitable for irrigation only in well-drained soils using excess
water to wash the soil and use salt tolerant crops.
C4 High salinity water in many cases is not suitable for irrigation. Must be used only
in highly permeable soils with good drainage using excess water to wash the soil
salts and using very salt tolerant crops.

C5 Excessive salinity water that should be used only in very rare cases, taking every
precaution.
C6 Water with excessive salinity, not suitable for irrigation.
S1 Water with low sodium content, suitable for irrigation in most cases. Problems
can occur in very sodium sensitive crops.
S2 Water with average sodium content, with a certain sodium accumulation danger
in the soil, especially in fine-textured (clay, clay loam, silty clay) of low
permeability. Should be monitored sodium level in the exchange complex and the
physical properties of soil, applying the necessary corrections.

S3 Water with high sodium content and high risk of sodium accumulation in soil.
Applications of organic matter and gypsum will be needed as a soil improver to
reduce sodium excess. It will require excellent drainage for washing.
S4 Water with high sodium content. In general is not suitable for irrigation, except in
soils with very low salinity and taking all recommended precautions.
Table 60 Diagram Interpretation of Riverside rules

The Riverside classification that includes the EC and RAS, measures the water hazard
separating the effect of salts in general and the sodium salts. The recommendations are
based on gravity irrigation, therefore caution should be applied when water classified with
this standard, is used in different forms of irrigation.

3. Content of toxic elements for plants. Elements in solution, that independently to the
above parameters, because of their concentration, are toxic.
1. Boron. Despite being an essential element for plants, at high concentrations
produce toxicities initially in the older leaves.
 131

-1
Water type
Boron levels (mg L ) according to crop
Sensible Semitolerant Tolerant
1 < 0.33 < 0.67 < 1.00
2 0.33 – 0.67 0.67 – 1.33 1.00 – 2.00
3 0.67 – 1.00 1.33 – 2.00 2.00 – 3.00
4 1.00 – 1.25 2.00 – 2.50 3.00 – 3.75
5 > 1.25 > 2.50 > 3.75
Table 61 Clasificación de las aguas por su nivel de B y tolerancia de cultivos.

TOLERANT SEMITOLERANT SENSITIVE

4 ppm 2 ppm 1 ppm

ASPARAGUS POTATOE WALNUT
DATE PALM COTTON WALNUT TREE
SUGAR BEET TOMATO NAVY BEAN
FODDER BEET RADISH PLUM TREE
BEETROOT PEA PEAR TREE
ALFALFA OLIVE APPLE TREE
BEAN BARLEY VID
ONION WHEAT KADOTA FIG
TURNIP CORN CHERRY TREE
CABBAGE SORGHUM PEACH
LETTUCE OATS APRICOT
CARROT PUMPKIN BLACKBERRY
PEPPER ORANGE TREE
SWEET POTATO AVOCADO
LIME BEAN GRAPEFRUIT
LEMON
0 3 PPM

Table 62 Relative crop tolerance to the presence of boron in irrigation water..

2. Chlorine. Ion that dissolved in water at high concentrations, generates toxic

effects, especially because it is an ion that is not retained in soil, and enters the
plant through the mass flow and is initially accumulated in the leaf tips, if the
concentration exceeds crops tolerance, burns initially be presented at the tips,
continuing along the edges until defoliation. This occurs when the Cl content in
leaves is 0.3 to 1.0% on dry weight.
 132

Class Chlorides (me/l)

Good Less than 1.0

Conditioned From 1.0 to 5.0

Not Recommended More than 5.0

Table 63 Classification of irrigation waters according to their content of chlorine

b) Sodium, Na. In addition to the direct damage that sodium causes soil structural
stability, also causes direct antagonists effects because it competes with other ions
such as potassium while taking nutrients, and generates toxic effects to accumulate
in the leaves of some crops according to its tolerance. When the waters with high
sodium levels are used for sprinkler irrigation or agrochemicals application, sodium
has a direct toxic effect on the leaf.

Na+ and Cl- Concentration in me/l

<5 5-10 10-20 >20
Almond Vid Alfalfa Cauliflower
Apricot Chili Barley Cotton
Citrus Potato Corn Beetroot
Plum Tomato Cucumber Sunflower
Sunflower
Table 64 Water Effect of Na and Cl when applied by sprinkler irrigation (Mass, 1984)

Use Restrictiveness
Potential Problem Units Mild to
None Severe
moderate
Salinity (affects water availability for crops) 2

ECa dS/m < 0.7 0.7 – 3.0 > 3.0

(o)
STD mg/l < 450 450 – 2000 > 2000
Infiltration (influence on the water infiltration rate into the soil.
Evaluate using CEa and RAS together) 3
RAS = 0 – 3 Y Cea = > 0.7 0.7 – 0.2 < 0.2
=3–6 = > 1.2 1.2 – 0.3 < 0.3
= 6 – 12 = > 1.9 1.9 – 0.5 < 0.5
= 12 – 20 = > 2.9 2.9 – 1.3 < 1.3
= 20 – 40 = > 5.0 5.0 – 2.9 < 2.9
Specific ions toxicity (influence on sensitive crops)
 133

Sodium (Na)4
Gravity irrigation RAS < 3 3–9 >9
Sprinkling irrigation me/l < 3 >3
Chlorine (Cl)4
Gravity irrigation me/l < 4 4 – 10 > 10
Sprinkling irrigation me/l < 3 >3
Boron (B)5 mg/l < 0.7 0.7 – 3.0 > 3.0
Miscellaneous Effect (Influence on susceptible crops)
Nitrgen (NO3 - N)6 mg/l < 5 5 – 30 > 30
Bicarbonate (HCO3)
(Foliar sprinkling only) me/l < 1.5 1.5 – 8.5 > 8.5
pH Normal Range 6.5 – 8.4
Table 65 Guidelines for interpreting the water quality and its restriction degree for agricultural irrigation.

1 Adapted from the consultants committee of the University of California, 1974.

2 ECa is the electrical conductivity in dS/m at 25°C or in millimhos units per centimeter (mmho/cm). Both are equivalent. STD
means total dissolved solids, reported in milligrams per liter (mg/l).

3 RAS is the sodium adsorption ratio. RAS is sometimes expressed as RNa. At a certain RAS, the infiltration rate increases as
water salinity increases. Evaluate the potential for infiltration problem in RAS amended by CEa.

4 Most crops and woody plants are sensitive to sodium and chlorine. Most annual crops are not sensitive. In the case of
sprinkling on foliage at relative humidities below 30% sodium and chlorine can be absorbed by the leaves and produce
toxicities in sensitive crops.

Plant nutrition

The nutrients

Until now, in plants we have found more than 70 chemical elements as part of its mineral
composition, some at higher rates than others, however, by the fact of being present in the
plant, does not mean they have a role in metabolism of this, many of them are present due
to the soil mineralogical composition.

Because plants do not have a quite selective absorption mechanism, can absorb both
essential and non-essential items and also harmful, so the plants mineral composition is not
a good guide to determine if the absorbed elements play or not an essential role in their
life. Techniques such as missing element can help determine its essence.

For normal growth of the plants 17 elements are considered as essential, also called
nutrients, which must meet certain requirements.
 134

Essentiality criteria.

In 1939, Arnon and Stout state that nutrients are those that meet the "Essentiality criteria"
listed below.

1. In the absence of the element in question cannot be a normal plant development

and this is unable to complete its life cycle.

2. The absence of an essential component leads to the appearance of deficiency

symptoms that could be solved only by the element in question, ie the element in
question may not be replaced or completely replaced by another element.

3. The function element within the plant metabolism must be known.

4. Its action must be direct on the plant, and not through the substrate favorable
modifications.

According to these researchers, the chemical elements that meet these criteria may be
considered as essential for the plants growth and development and therefore are called
nutrients.

These criteria clearly conclude that the lack of a nutrient results in damage, abnormal
development or death of the plant.

Nutrient classification.

1) According to their plant requirement level

Classification Plant requirement
Necessary Those vital for plant nutrition importance and that meet the
elements requirements stipulated above, in essentiality criteria: C, H, O, N, S, P, K,
Ca, Mg, Fe, Mn, Cu, Zn, B, Mo, Cl, y Ni.
Useful elements Elements that benefit the nutrition of higher plants or specific groups of
them, without being necessary: Si, Co y Na.
Expendable Elements that are absorbed by plants, but not perform specific
elements physiological functions either directly or indirectly benefit the growth of
plants.
 135

2) According to the amounts required by the plants.

Macronutrients: its concentration in plant tissue is may 1000 mg.kg -1(ppm
Micronutrients: Its concentration in plant tissue is less than 500 mg.kg-1(ppm)

ELEMENT SYMBOL ABSORPTION FORM

MACRONUTRIENTS
Carbon C CO2
Hydrogen H H2O
Oxygen O H2O, O2

Nitrogen N NH4+, NO3-

Phosphorus P H2PO4-, H2PO42-
Potassium K K+
Calcium Ca Ca2+
Magnesium Mg Mg2+
Sulfur S SO42-
Table 66 macro-nutrients to plants and their way of absorption. Bennet, 1993.

ELEMENT SYMBOL ABSORPTION FORM

MICRONUTRIENTS

Iron Fe Fe2+, Fe3+

Zinc Zn Zn2+, Zn (OH)20
Manganese Mn Mn2+
Copper Cu Cu2+
Boron B B(OH)30
Molybdenum Mo MoO42-
Chlorine Cl Cl-
Silica Si Si(OH)40
Sodium Na Na+
 136

Cobalt Co Co2+
Vanadium V V+
Nickel Ni Ni2+
Table 67 Essential macro-nutrients to plants and their way of absorption. Bennet, 1993.

3) According to their absorption way (anions or cations). The pH of the rhizosphere region
may vary up to two units or more compared to the soil pH and this has effects such as:

a. Absorption imbalance in ratio cation/anion.

b. The corresponding difference that generates the above, regardin H+ y HCO3-
(u OH-) excretion and
c. Organic acid excretion
The supply form of N influences the absorption ratio cation/anion and the
rhizosphere pH. Nitrates supply associated with a high excretion of bicarbonate
more than with hydrogen excretion, while the opposite happens with ammonia. In
alkaline or neutral acidification rhizosphere of fed ammonium plants can promote
the mobilization of soluble calcium phosphates and, therefore, phosphorus
absorption and micronutrients absorption as boron, iron, manganese and zinc.

CATIONS ANIONS UNDISSOCIATED MOLECULES

NH4+ NO3-
6K+ H2PO4-
Ca2+ SO42-
Mg2+ Cl-
Fe2+ MoO42-
Mn2+ H3BO3
Zn2+
Cu2+
Ni2+
Table 68 Common ionic forms in which nutrients are absorbed by plants.

4) According to their chemical nature and their physiological functions. The element
requirement from the physiological point of view is not related to the present element
amount in the plant because the plant can absorb greater than necessary amounts in a
given time or the physiological need for an item will be determined by other element
absorbed amount, like in Mo case regarding absorbed forms of N, therefore it is difficult
to establish the nutrient need in an absolute amount for a given
 137

plant, also affected by antagonism and synergism interactions between nutrients,

the specific element physiological role and its absorption influence. So this is why it
the classification becomes important, as a macronutrient may behave in a similar
way to a micronutrient and this cannot be identified in other classification

Nutrient Chemical Absorption Physiological function

Group

C, H, O, Nonmetallic As CO2, HCO3-, H2O, NO3- Major constituents of

N, S. , NH4+, N2, SO42-, SO2. biomolecules. Participate in all
The ions are taken from enzymatic reactions and
the soil solution and metabolic processes; protein
gases from the synthesis, breathing and
atmosphere. photosynthesis.

P, B, Si Nonmetallic Oxo complex shaped; Participate in esterification

phosphates, borates reactions with alcohol groups
(boric acid), silicates (hydroxyl) of numerous
from soil solution. molecules. The phosphate esters
are responsible for the main
energy transfer reactions.

K, Na, Ca, Alkali and In cation form in soil As enzyme activators modify the
Mg. alkaline earth solution. protein conformation (tertiary
Metals structure). Involved in the
regulation of osmotic potential
and anion balance controlling
membrane permeability and
electropotentials.

Fe, Mn, Heavy Metals In cation form, ions or Become chelates and are
Cu, Zn, chelates of soil solution. incorporated into enzyme
Mo, Ni, prosthetic groups. Involved in
Co electrons transport through
valence changes.

Cl Halogens Chloride form of the soil Esencial en la fotolisis del agua en

solution and through the el fotosistema II y en el balance de
foliage in coastal areas. cationes. Essential in water
 138

photolysis in photosystem II and

in cations balance.

Table 69 Nutrients classification and beneficial elements according to their chemical properties and physiological
functions (Mengel and Kirkby modified 1987). B and Mo are ttransition metals or metalloids with predominance of
nonmetals. Na and Co are beneficial and essential for some species.

5 ) By specific function

STRUCTURAL ENZYM CONSTITUENT OSMOTIC TRANSPORT AND

REGULATION

The element is part of one
molecule or more organic
compounds.
Special case of the first The nutrient form low-
group, transition or energy bonds with organic
elements metals (Mo). Are molecules of low molecular
parts of the prosthetic weight, to facilitate their
enzymes group and are mobility from one organ to
essential for the activity another. It is stored in
thereof. vacuole for osmoregulation
processes.

N: aminoacids and proteins. Mo, Cu, Fe, Mn, Zn, Ni, Potassium fulfills these two
participate as cofactors functions, sodium in some
Ca: Middle lamella pectate participan como cofactors cases serves as
of the cell wall. (activators) of several osmoregulatory.
Mg: Occupies the center of enzymatic reactions.
the tetrapyrrole core of the
chlorophyll molecule.

P: as an integral part of
nucleotids and nucleic acids.

An element can perform several functions corresponding to one or various vital processes
such as photosynthesis, breathing, amino acids or proteins synthesis, transfer of hereditary
traits, and others.
 139

STRUCTURAL ENZYME
Table 70 ENZYME Functions that
can be CONSTITUENT ACTIVATOR performed by a
structural element.

6) For its mobility: Indicates the ease with which an element is recycled from old to
new leaves, and helps identify their deficiencies with this mobility pattern. The mobile
elements will initially present their deficiency in older leaves and immobile elements
initially present their deficiency in new leaves.

Movable Mid mobility Immobile

N, P, K, Mg, Ni S, Cu, Zn, Mo, Cl, Na Ca, Mn, B, Fe

Table 71 Elements and its mobility

7) Beneficial elements. Elements that stimulate growth but are not essential or are
essential but only for a certain group of plants. Also those elements that can offset the
toxic effects of others, as well as those that can replace the essential nutrients in a less
specific function, such as maintenance of osmotic potential. These are Na, Si, Co, Al, Se,
Ni, I, and Va.
 140

Nutrients functions

The function of essential elements is briefly described in this section, with the
understanding that a complete description of biochemical, physiological and ecological
functions would be excessive.

Nitrogen

This nutrient, sometimes regarded as the highest concentration (between the called
minerals) in plant tissue (4.0 - 6.0%), can be absorbed as much as ammonium nitrate. As
the more absorbed ammonium form and used by plants in their juvenile stages and nitrate
in their adult stages, although different plant species vary use proportion, depending on
environmental conditions for growth; eg rice almost always absorbs and uses ammonium.
This element covers many plant functions and is constituent many organic compounds as
amines, amides, sugars, pigments, aminoacids, proteins and enzymes, as well as
chromosomes. It is highly mobile within the plant, as their forms are translocated, usually
at the growing points (Westerman, 1990).

Phosphorous

It is an essential element of difficult dosage form for plants. Is required in considerably

smaller amounts than Nitrogen. It is present from 0.2 to 0.8%. Is absorbed in anionic forms,
usually as ortho primary phosphate ion (H2PO4-), depending on soil pH. In plant is present in
constitution compounds (phosphate sugars, nucleic acids, phospholipids and coenzymes)
and compound transporters of energy (ATP, NADP, FAD) (Westerman, 1990). Phosphorus is
transported in the xylem as inorganic phosphates and phosphorylcholine, and in phloem as
sugar phosphate and inorganic phosphate.

Its main physiological role is the phosphorylation of the organic substances. Phosphoric
acid is temporarily combined with a carbonyl, enolic or nitric group, to form an energy-rich
compound: adenosine triphosphate (ATP), which, decomposing into ADP, releases this
energy used in metabolic processes (Baeyens, 1970).

Potassium

Is an important part of tissue (1.5 - 6.0%), absorbed as a cation (K+); within the plant is the
cytoplasm and helps keep turgid. It usually focuses on the growing points. It is an easy
 141

element leaching, since in each irrigation, washing amounts are significant (Westerman,
1990).

K also plays an important role in maintaining membrane stability and of such absorption
capacity to generate relevant osmotic and hydric potentials. (Baeyens, 1970).

Unlike Nitrogen, Phosphorus, Sulfur and various other; Potassium is not an integral part of
components of the plant as the protoplasm, cellulose and fat, its function seems rather
catalytic nature. Despite this, it is imperative for the following physiological functions:

1. For the synthesis of carbohydrates or starch formation and transformation, also in

manufacturing albumin.

2. Participate in Nitrogen metabolism and proteins synthesis.

3. Controls and regulates the activities of several essential minerals.

4. Participate in the neutralization of physiologically important organic acids. These acids

tend to lower the pH of the cell sap, which are neutralized by K and, curiously, by the Na.
Substances called buffers, consists of weak acids combined to an alkaline or alkaline earth
cation. These cations are represented up to 90% by K and only 10% for Na, Ca, Mg and Al.
This highlights the importance of K versus other cations (Baeyens, 1970).

5. It is an activator of several enzymes, which catalyze phosphorylation and conversely

inhibits breathing.

6. Promotes growth in meristematic tissues.

7. Adjust the stomatal opening and participates in creating water potential gradients
between tissues, regulating the flow of water between them.

Calcium

Is an element of structural constitution in the plant, its concentration varies from 0.4 to 2.0%, is
part of the cell walls in the form of calcium pectate, which plays an important role in the
division thereof and in cell elongation. It is also required to achieve integrity and membrane
stability. It is part of the enzyme α-amylase. Usually precipitate vacuoles as calcium oxalate. It
has also been shown that Calcium favors the formation and increasing the protein contained in
mitochondria. If so, the role played by mitochondria in aerobic respiration, and
 142

hence in the assimilation of salts, indicates that there must be a direct relationship
between the Calcium ion and assimilation in general. Calcium is translocated in ionic form
positive divalent (Ca2+). This is a relatively immobile element in the plant. Almost does not
move in the phloem (Westerman, 1990).

Magnesium

Mg concentration in plant tissue varies from 0.2 - 1.0%. It is absorbed in divalent positive
ionic form (Mg2+). This element intervenes in many enzymatic reactions (as activator) and
in the assimilation of Phosphorus and forms the core of the chlorophyll molecule. It is
highly mobile in the plant and is required for the integrity of the ribosome structure
(Westerman, 1990).

Magnesium is required for the activation of many enzymes involved in the metabolism of
carbohydrates and is very important in the citric acid cycle, of importance in cellular
breathing. Numerous phosphorylation reactions are related to nitrogen metabolism in the
plant are catalyzed by this element.

Sulfur

Is part of various organic compounds, among which highlits some aminoacids like cysteine
and methionine, which are essential for the protein formation. Intervenes in the synthesis
of vitamins and phytohormones and is part of thiamine, biotin. It is present in the
glycosides that give the characteristic smell of garlic, onions and others (Tisdale et al.,
1985). Form some other compounds as sulfolipids that are an important part of
membranes. Is a structural constituent of certain coenzymes (Coenzyme A) and prosthetic
groups. It is also involved in redox reactions (Marschner, 1986).

Iron

Iron plays an important role by being activator of the enzyme and essential system for the
formation of chlorophyll precursors and electron transfer in various oxide reduction
reactions. It is present in the cytochromes, which act as electron carriers in photosynthesis
and plants breathing. It is an essential part of the ferredoxin and probably of nitrogenase
and nitrate reductase and other enzymes activator.
 143

Manganese

Is absorbed in the form of manganese ion (Mn2+) and in molecular combination with certain
organic complexes such as EDTA. Availability depends on the soil reaction (pH). It is a trigger for
various enzyme systems associated with carbohydrate and fatty acids metabolism,
phosphorylation reactions and nucleic acids formation (DNA, RNA) and also isocitrate
dehydrogenase enzyme activator in the Krebs cycle, in conjunction with other metals in
enzymes activation such as arginase and phosphatase. It is involved in the water photolysis in
photosynthesis, as well as in the sucrose synthesis in the plant. Forms metalloproteins.

Zinc

Zinc is absorbed by the plants roots in the form of Zn2+ ion and can also be absorbed in
chelate form as EDTA. This element is involved in the certain amino acids synthesis. The
most important role of zinc in plants is the enzymes metal activator among which enolase,
aldose, oxaloacetic decarboxylase, alcohol dehydrogenase, lactic acid dehydrogenase,
carboxypeptidases, and many other, among which stands out the auxin synthesis (indole
acetic acid).

Copper

Copper is absorbed by plants in cupric ion (Cu2+) form, and can also be absorbed as an organic
complex salt such as EDTA. Copper is a metal activator of several enzymes among which are
tyrosinase, laccase, ascorbic acid oxidase and dehydrogenase butyryl Co-A. Is also said that
could be one of the related metals in the light reaction in plants. Is an electron carrier in
photosynthesis as part of plastocyanin. It is also part of the polyphenol and nitrate reductase.

Molybdenum

Is probably absorbed as (MoO42-) ion. It is part of the plant enzymatic reactions, acting as a
cofactor (Westerman, 1990) in the reduction enzymes of nitrate (NO3) to ammonium (NH4)
and N2 biological fixation in legumes.

Nitrogen fixation and reduction are, of course, prior requirements to the aminoacid and
protein synthesis, and a molybdenum deficiency has adverse effects on them. Molybdenum
 144

is also involved in nitrate accumulation in an apparent decrease in ascorbic acid oxidase

activity. Molybdenum is known to be specific for the nitrate reductase enzymes and
xanthine oxidase activation.

Boron

It is involved in the sugar and starches transport through the membrane and into the
phloem conduits. Is also involved in cell wall synthesis and carbohydrates metabolism;
interact with auxins (Gupta, 1979). Boron also influences cell growth and elongation;
affects carbohydrate metabolism and plays an essential role in the aminoacids formation
and protein synthesis (Gupta, 1979; Tisdale et al., 1985).

Chlorine

Take part in the capture and accumulation of light energy processes, through its
intervention in the water photolysis reactions and phosphorylation in photosynthesis.
Participates with K as osmoregulator and as anion in anion - cation balance in plants.

Carbon, Oxygen e Hydrogen

Are part of all basic structures and chains of all organic molecules of vegetal metabolism.
These 3 nutrients make up 85-96% of the elements present in plants.

Silica

It is found in the cell membrane and the epidermis at the perspiration organs periphery and
along the vascular vessels. Its function is to stiffen these structures. No physiologically
active, but protects the peripheral tissues and various fungal and pest attacks. Some
evidence suggests a role in roots development (Baeyens, 1970).

Vanadium

It is necessary for some types of algae. Their addition favors the activity in plants of ascorbic
acid, peroxidase and catalase activity, in leaves and fruits. It also stimulates the invertase
activity. (Baeyens, 1970). Its essentiality in some higher plants is not fully demonstrated.
 145

ELEMENT ABSORPTION CONCENTRATION MAIN FUNCTIONS

FORM (%)

Macronutrients:

Carbon CO2  44 % Organic compounds component.

Oxygen H2O u O2  44 % Organic compounds component.

Hydrogen H2O 6% Organic compounds component.

Nitrogen NO3- o NH4+ 1-4 % Aminoacids, proteins, nucleotides,

nucleic acids, coenzymes and
chlorophyll.

Potassium K+ 0,5-6 % Enzymes, amino acids, and protein

synthesis.
Activator of many enzymes.
Stomata opening and closing.

Calcium Ca2+ 0,2-3,5 % Calcium from cell walls.

Enzym cofactor.
Cell permeability.
Calmodulin component, a regulating
membrane and enzyme activities.

Phosphorus H2PO2- o 0,1-0,8 % Formation of "high energy"

HPO42- phosphate compounds (ATP and
ADP).
Nucleic Acids.
Sugars phosphorylation.
Several essential coenzymes.
Phospholipids.

Magnesium Mg2+ 0,1-0,8 % Part of the chlorophyll molecule.

Activator of many enzymes.

Sulfur SO42- 0,05-1 % Some amino acids and proteins.

Coenzyme A.
 146

Micronutrients:

Iron Fe2+ o Fe3+ 25-300 ppm Synthesis of chlorophyll,

cytochromes and nitrogenase.

Chlorine Cl- 100-10.000 ppm Osmosis and ion balance, probably

essential in photosynthetic
reactions that produce oxygen.

Copper Cu2+ 4-30 ppm Activator of many enzymes.

Manganese Mn2+ 15-800 ppm Activator of many enzymes.

Zinc Zn2+ 15-100 ppm Activator of many enzymes.

Molybdenum MoO42- 0,1-5,9 ppm Nitrogen fixation. Nitrate reduction.

Boron BO3- o B4O72- 5-75 ppm Influence in calcium utilization.

Essentials elements for some plants or organisms:

Cobalt Co2+ Traces Required by nitrogen-fixing

microorganisms.

Sodium Na+ Traces Osmotic and ionic balance, not

essential for many plants. Required
by some desert species and
wetlands. It may be necessary in all
plants that use C4 photosynthesis.

Figure 63 Summary of the most important functions of inorganic nutrients in plants, Taiz, L. And Zeiger, E., 1988.

N P K Ca Mg Na Fe Cu Mn B Mo Zn
DEFICIENCIES SERIES
Without N.................... -- -- ++ ++ (+) (+) (+) (+) (+) (++) + (+) 0
Without P.................... ++ -- -- (++) + (+) (--) (+) (+) (++) + (+) (+)
Without K.................... ++ ++ -- -- + ++ ++ (+) ++ (++) (+) (++) ++
Without Ca.................. + + (+) -- -- 0 + (--) 0 (+) (+) (-- --) ++
Without Mg................. 0 + (+) + -- -- + 0 0 0 (+) (+) 0

Without Fe.................. 0 0 (--) 0 0 (--) (--) + 0 0 -- 0

 147

Without Cu.................. -- + + + (+) (+) + -- 0 + + +

Without Mn................. 0 0 0 0 0 0 0 + -- 0 + 0
Without B................... 0 (+) (+) (+) 0 0 + 0 + -- -- 0 0
Without Mo................. 0 0 0 0 (+) 0 -- 0 0 0 -- -- --
Without Zn.................. 0 0 (+) 0 0 (--) 0 + 0 -- + --
EXCESS SERIES:
More N ................. ++ (--) (--) 0 ( --) (+) -- (--) 0 0 -- -- 0
More P.................. 0 ++ (+) -- -- ++ -- 0 (--) 0 (--) (--)
More K.................. -- -- + -- -- -- (--) 0 (--) 0 (--) 0
More Ca................ -- -- (--) + -- -- 0 (--) -- -- ++ 0
More Mg............... 0 0 (+) -- + 0 (--) (+) 0 0 0 (+)

More Fe................ + 0 0 + (--) + (+) 0 (+) 0 + 0

More Cu................ 0 0 0 0 0 0 -- ++ 0 0 0 0
More Mn............... 0 0 0 0 -- 0 -- 0 + 0 -- 0
More B.................. 0 0 0 0 -- 0 0 0 0 ++ + --
More Mo............... 0 0 0 0 0 0 0 0 0 0 ++ 0
More Zn................ 0 0 0 0 0 0 -- 0 0 0 0 0
Figure 64 Relationship of synergism and antagonism between different nutrients. Penningsfeld y kurzmann(1975). ).

NOTA:+ increase and – decrease in the plant content (dry weight). O indicates no difference with the control.
The double signs ++ and - - indicate differences in comparison with the control of a minimum content of 50%
of macroelements and 30% of microelements; the sign ( ) indicates that the deviation is not present in all
plants tested in the same direction.

Keys for the identification of nutrient deficiencies

1.- Color changes in old or lower leaves, the most relevant, localized or generalized symptoms; alteration in the overall
plant growth.

1.1.- Symptoms spread throughout the plant 1.1.- Symptoms spread throughout the plant

1.1.1.- Nitrogen. Chlorosis throughout the plant; older leaves turn yellow from the tip to the base,
dehydrated with pale brown in a "V" form, (maize and sorghum). Short and slender stems;
tillering in cereals is low. The root relative to the aerial part is very long and with few branches.
The plants are small and spun with rigid appearance. The fruits are also small.

1.1.2.- Phosphorus. Dark green foliage with purple tones in leaves, petioles, and veins. Short and
slender stems, poor tillering in cereals. Root slightly branched long reddish brown blotchy. The
plants are small like their leaves, seeds and fruits with rigid appearance. In the presence of cold
damage, purple hues are also developed.
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1.2.- Symptoms mostly localized 1.2.- Symptoms mostly localized

1.2.1.- Potassium. Coffee and necrotic discoloration in old leaves from the tips and moving
towards the base margins (maize and sorghum), the cotton sheets are frequently thick and brittle
(stilted). The stems are weak and plants lodge. The roots are long, yellowish and "leathery" with
little branching, plants appear to wilt, and has little resistance to cold. The fruits and seeds are of
poor quality.

1.2.2.- Magnesium. Interveinal chlorosis in older leaves of dicotyledonous plants with cotton
purple red or red wine stripes. In grasses chlorotic points necklace is observed along the veins
that may be surrounded by a strong redness. Usually thin stems and leaves.

2.- Symptoms in new upper leaves and eventually intermediate sheets

2.1.- The terminal bud grows normally. 2.1.- The terminal bud grows normally.

2.1.1.- new leaves not wilted, chlorosis but without necrosis in basal zone or distributed in whole
leaf part.

2.1.1.1.- Manganese. Leaves grayish-yellow or reddish gray with green veins, soon comes the
necrosis, dead areas will fall and give the plant a frayed look. In oats causes gray sins ("gray
speck"). Deficiency often occurs in middle leaves. Estate underdeveloped and brownish. In
grasses chlorosis occurs in stripes and dicotyledonous appears a mosaic.

New leaves show chlorosis usually without necrosis; veins may be pale or dark and partly chlorotic.

Iron. Chlorosis of new leaves, plants and margins retain color longer. The ribs remain green and
in extreme cases are also chlorotic. The affected sheet is bent upwards. Eventually the entire leaf
turns yellow-whitish. The roots are short brown and profusely branched. It occurs in alkaline
soils, but also in manganiferous ground.

2.1.1.2.2.- Sulfur. New leaves greenish yellow including the ribs. Often paler than the leaf blade
veins. The plant is generally pale green and gives similar to the lack of nitrogen patterned
appearance. The stems are slender, woody and abnormally with elongated internodes. The root
is profuse and white.

2.1.2.- New leaves no withered

2.1.2.1.- Copper. In grass leaves are rolled and turn white without turn yellow. Strong tillering,
thin stems. It inhibits or disrupts the formation of spikes. White and empty spikes. In dicots new
leaves are hung and sometimes have bad form. The generative growth is more affected than the
vegetative. It makes a bad form and affects the inflorescence. In fruit the ends of the branches
are resected and defoliate. The root growth is limited and rootlets are long thin and white.
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2.2.- The terminal bud dies after new leaves are bad 2.2.- The terminal bud dies after new
formed from the tip or base.
leaves are bad formed from the tip or base.

2.2.1.- Calcium. The new leaves of the tips of the branches are twisted. The inflorescences are
dried as blight damage. Apical rot is typical in tomato, pepper, cucumber, watermelon. Cause
numbness in apple fruit. In lettuce and other vegetables marginal necrosis develops. The roots
are short and dark brown or black.

2.2.2.- Boron. Thick, small and deformed buds. The new leaves of the terminal bud fade from its
base. The sheet is deformed from its base. The sheet is deformed and twisted with a cardboard
rough appearance. The internodes are short, growth stops and growing tips die. There are gaps
or cracked stems and in tubers there is coffee heart. The roots are very short and thick with
rounded tips brown.

3.- Symptoms in new or old leaves

3.1.- Molybdenum. Symptoms are similar to the lack of nitrogen in legumes. In tomato and citrus
appears yellowish mottling. The margins and the tips become necrotic as in nitrates excess.
Intervenal perforations. The core sheet grows but not the leaf blade. In cauliflower tail whip is
given. Are often more affected new leaves.

3.2.- Zinc. Chlorosis progresses rapidly with dead zones in new or old leaves. (Chlorosis can reach
the white color). The new leaves are very small, narrow and thick. The stems are very short
internodes, "dwarfism". Often there is rosetting on the tips of the growing shoots. The root
development is limited a little.

Table 72 Codes of deficiency identification, Uvalle B.

The laws of plant nutrition.

Assuming that plant nutrition should not be a limiting factor in agricultural production and
that the optimum plants supply with nutrients in each of the stages of growth strongly
influences the high performance and good quality of the harvest, have developed a set of
principles that relates the plant nutrition to performance, resulting laws. Not to mention, as
has been said above, that proper crop development is the result of the interaction of
several factors that have been linked previously.

1. The Liebig's law of the minimum. Established in 1855. The crop yield is governed by the
most limiting factor, and that only correcting that limiting can increase the performance
factor. When this factor is corrected, yields become regulated by the following limiting
factor. Further increases will occur only if that factor is corrected. This process is
repeated with performance-stepped increments until no limiting factors exist. This law
requires balanced elements contributions of fertilization, so that none is limiting. So, we
will have to look A BALANCE IN THE FERTILIZATION, even
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more than the absolute value of the fertilization amounts. Any factor that affects
the plant development can be limiting if is not within the required parameters.

Example of limiting factors of performance, ‘’Minimum barrel’’ The nutrient in

less quantity limits performance, here N and Mg respectively.
Limitin factors
NOWADAYS (Frequent
PREVIOUSLY secondary elements or
(Frequent N) trace elements

Wheat yield
ton/ha

TRACE ELEMENTS
Figure 2 Ley del mínimo de Liebig.

2. Optimum law. The minimum factor most strongly influences the performance, the
more they are in optimal the other factors. Mayer, 1869; Wollny, 1877; Liebig, 1895.
3. Law of diminishing returns, Misterlich, 1909. The increases in performance are
decreasing as growth factor increases, also indicates that performance is influenced
by all limiting factors simultaneously. The influence of each of the limiting factors is
proportional to the limitation degree. Misterlich was the first to develop an
equation that expresses:
a) = C ∗ (A − Y)

Where: dx=Increasead growth factor

dy= Increase in growth due to dx
C= Proportionality constant
A= Maximum possible performance
Y= Yield before applying dx
Integrating the equation and considering that y=0 when x=0. We have the
following representation.
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Growth Increasead
factor units Harvest % harvest (%)

0 0 0

1 50 50

2 75 25

3 87.5 12.5

4 93.75 6.25

5 96.88 3.13

6 98.44 1.56

7 99.22 0.78

8 99.61 0.39

9 99.8 0.19

10 99.9 0.1

Table 73 Representation of Misterlich equiation, Baule units.

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MISTERLICH SCALE REPRESENTATION, BAULE UNITS

120

100
MAXIMUM YIELD %

80

60

Harvest %
40

Increase in
20 Harvest %

0
0 1 2 3 4 5 6 7 8 9 10

GROWTH FACTOR UNITS

Figure 66 Misterlich scale representation.

Baule Unit: Term for the amount of nutrients that is associated with 50% of maximum yield.
If 50% of maximum yield is obtained with 1 Baule unit of a nutrient (Example 5 kg S/ha),
then 75% of the yield is associated with 2 Baule units, 87.5% of maximum yield with 3 Baule
units and so in turn.

An example of the interaction of two factors is: if you know that there are 2 N Baule units
and 3 P Baule units of in the soil, you'd think that there is N required to achieve 75% of
maximum yield, and sufficient P to reach 87.5% of maximum yield. Therefore, one would
estimate that would be reached at least 75% of the production considering the narrower
element. In fact due to the interaction of nutrients, one will get only 66% of maximum
performance, since it has little N and P. The result is obtained by multiplying 75% of N by
87.5% of P.

To properly implement this law we should consider the following:

a) Knowing the soil reserve level to each of the nutrient elements.

b) Keep in mind the fertility level in relation with the considered element threshold.
c) With microelements, consider that performance is achieved quickly but then begins
a depressive effect.
d) Water has a very low efficiency, so it is applied in large quantities.
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4. Law of Maximum, Voisin, 1945. Excess of one or more nutrients has a decreasing
effect in performance.

Appropiate range

Luxury consumption

Deficiency range Toxicity range

Visible symptoms
deficiency
G rowth or perf orma nce

Nutrient contents in plants tissue

Figure 67 Nutrient sufficiency ranges and its effect on performance.

5. Restitution Law, Boussignault, 1860. “If you want to maintain soil fertility is essential
to restore the nutrient elements extracted in harvest.” This is a necessary but not
sufficient law to proper feed with minerals the crops, since:
a. There are several extraction crop losses that are necessary to restore.
b. Does not include future increases in crop yields.

6. Extended restitution Law. “To maintain soil fertility is necessary to restore all the
nutrients that may be lost for any reason” This Law is only for the maintenance and
preservation of the soil fertility current status. From the fertilizer calculus point of
view, the soils can be classified as poor, medium or rich:
a. Rich soils: There is no response to fertilizer application.
b. Medium soils: The responses positive or negative will depend on factors
other than nutrition, and in this kind of soils applies a 100% the
“conservation compost”
c. Poor soils: The crop answer is always positive to fertilization and its
necessary applies correction compost.
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After this concepts review is clearly noted that plant nutrition not only refers to fertilizers
application, as such, indiscriminately, fertilization with a certain element is perhaps the
final practice in a crop which must be observed and change in the extent possible, the
factors that will make optimal photosynthesis and minimum respiration, in any case using
compounds that support the plant best performance (photosynthesis-respiration), in each
of its phenological stages and with nutritional relationships management direct the plant
behavior towards the component performance of economic importance development,
which is why the plant is being grown.

Grown in a soil with good physical and chemical characteristics, irrigate water with good
quality and cultivate under proper weather, whether if they are natural or modified, will
maximize plant performance and reduce own energy costs by optimizing its productivity.

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