1

Plant nutrition is the study of the chemical elements that are necessary for growth. In 1972, E.
Epstein defined 2 criteria for an element to be essential for plant growth: (1) in its absence the
plant is unable to complete a normal life cycle or (2) that the element is part of some essential
plant constituent or metabolite, this is all in accordance with Liebig's law of the minimum.[1]
There are 17 essential plant nutrients. Carbon and oxygen are absorbed from the air, while other
nutrients including water are obtained from the soil. Plants must obtain the following mineral
nutrients from the growing media:[2]

 the primary macronutrients: nitrogen (N), phosphorus (P), potassium (K)

 the three secondary macronutrients such as calcium (Ca), sulphur (S), magnesium (Mg).
 the macronutrient Silicon (Si)
 and micronutrients or trace minerals: boron (B), chlorine (Cl), manganese (Mn), iron
(Fe), zinc (Zn), copper (Cu), molybdenum (Mo), nickel (Ni), selenium (Se), and sodium
(Na).

The macronutrients are consumed in larger quantities and are present in plant tissue in quantities
from 0.2% to 4.0% (on a dry matter weight basis). Micronutrients are present in plant tissue in
quantities measured in parts per million, ranging from 5 to 200 ppm, or less than 0.02% dry
weight.[3]

Most soil conditions across the world can provide plants with adequate nutrition and do not
require fertilizer for a complete life cycle. However, man can artificially modify soil through the
addition of fertilizer to promote vigorous growth and increase yield. The plants are able to obtain
their required nutrients from the fertilizer added to the soil. A colloidal carbonaceous residue,
known as humus, can serve as a nutrient reservoir.[4] Besides lack of water and sunshine, nutrient
deficiency is a major growth limiting factor.

Nutrient uptake in the soil is achieved by cation exchange, where in root hairs pump hydrogen
ions (H+) into the soil through proton pumps. These hydrogen ions displace cations attached to
negatively charged soil particles so that the cations are available for uptake by the root.

Plant nutrition is a difficult subject to understand completely, partially because of the variation
between different plants and even between different species or individuals of a given clone. An
element present at a low level may cause deficiency symptoms, while the same element at a
higher level may cause toxicity. Further, deficiency of one element may present as symptoms of
toxicity from another element. An abundance of one nutrient may cause a deficiency of another
nutrient. Also a lowered availability of a given nutrient, such as SO2−4 can affect the uptake of
another nutrient, such as NO3–. Also, K+ uptake can be influenced by the amount NH4+ available.
[4]

The root, especially the root hair, is the most essential organ for the uptake of nutrients. The
structure and architecture of the root can alter the rate of nutrient uptake. Nutrient ions are
transported to the center of the root, the stele in order for the nutrients to reach the conducting
tissues, xylem and phloem.[4] The Casparian strip, a cell wall outside of the stele but within the
root, prevents passive flow of water and nutrients to help regulate the uptake of nutrients and
water.[4] Xylem moves water and inorganic molecules within the plant and phloem counts
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organic molecule transportation. Water potential plays a key role in a plants nutrient uptake. If
the water potential is more negative within the plant than the surrounding soils, the nutrients will
move from the more higher solute (soil) concentration to lower solute concentration (plant).

There are 3 fundamental ways plants uptake nutrients through the root: 1.) simple diffusion,
occurs when a nonpolar molecule, such as O2, CO2, and NH3 that follow a concentration gradient,
can passively move through the lipid bilayer membrane without the use of transport proteins. 2.)
facilitated diffusion, is the rapid movement of solutes or ions following a concentration gradient,
facilitated by transport proteins. 3.) Active transport, is the active transport of ions or molecules
against a concentration gradient that requires an energy source, usually ATP, to pump the ions or
molecules through the membrane.[4]

Nutrients are moved inside a plant to where they are most needed. For example, a plant will try
to supply more nutrients to its younger leaves than its older ones. So when nutrients are mobile,
the lack of nutrients is first visible on older leaves. However, not all nutrients are equally mobile.
When a less mobile nutrient is lacking, the younger leaves suffer because the nutrient does not
move up to them but stays lower in the older leaves. Nitrogen, phosphorus, and potassium are
mobile nutrients, while the others have varying degrees of mobility. This phenomenon is helpful
in determining what nutrients a plant may be lacking.

A symbiotic relationship may exist with 1.) Nitrogen-fixing bacteria, like rhizobia which are
involved with nitrogen fixation, and 2.) mycorrhiza, which help to create a larger root surface
area. Both of these mutualistic relationships enhance nutrient uptake.[4]

Though nitrogen is plentiful in the Earth's atmosphere, relatively few plants engage in nitrogen
fixation (conversion of atmospheric nitrogen to a biologically useful form). Most plants therefore
require nitrogen compounds to be present in the soil in which they grow. These can either be
supplied by decaying matter, nitrogen fixing bacteria, animal waste, or through the agricultural
application of purpose made fertilizers.

Hydroponics, is growing plants in a water-nutrient solution without the use of nutrient-rich soil.
It allows researchers and home gardeners to grow their plants in a controlled environment. The
most common solution, is the Hoagland Solution, developed by D. R. Hoagland in 1933, the
solution consists of all the essential nutrients in the correct proportions necessary for most plant
growth.[4] An aerator is used to prevent an anoxic event or hypoxia. Hypoxia can affect nutrient
uptake of a plant because without oxygen present, respiration becomes inhibited within the root
cells. The Nutrient film technique is a variation of hydroponic technique. The roots are not fully
submerged which allows for adequate aeration of the roots, while a "film" thin layer of nutrient
rich water is pumped through the system to provide nutrients and water to the plant.

[edit] Processes important to plant nutrition

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Plants uptake essential elements from the soil through their roots and from the air (mainly
consisting of carbon and oxygen) through their leaves.

In the leaves, transpiration occurs, where stomata open to take in carbon dioxide and expel
oxygen that drives the movement of water and nutrients to the plant[4] Green plants obtain their
carbohydrate supply from the carbon dioxide in the air by the process of photosynthesis.

[edit] Functions of nutrients

Each of these nutrients is used in a different place for a different essential function.[5]

[edit] Macro nutrients

Carbon

Carbon forms the backbone of many plants biomolecules, including starches and
cellulose. Carbon is fixed through photosynthesis from the carbon dioxide in the air and
is a part of the carbohydrates that store energy in the plant.

Hydrogen

Hydrogen also is necessary for building sugars and building the plant. It is obtained
almost entirely from water. Hydrogen ions are imperative for a proton gradient to help
drive the electron transport chain in photosynthesis and for respiration.[4]

Oxygen

Oxygen is necessary for cellular respiration. Cellular respiration is the process of

generating energy-rich adenosine triphosphate (ATP) via the consumption of sugars made
in photosynthesis. Plants produce oxygen gas during photosynthesis to produce glucose
but then require oxygen to undergo aerobic cellular respiration and break down this
glucose and produce ATP.

Phosphorus

Phosphorus is important in plant bioenergetics. As a component of ATP, phosphorus is

needed for the conversion of light energy to chemical energy (ATP) during
photosynthesis. Phosphorus can also be used to modify the activity of various enzymes
by phosphorylation, and can be used for cell signaling. Since ATP can be used for the
biosynthesis of many plant biomolecules, phosphorus is important for plant growth and
flower/seed formation. Phosphate esters make up DNA, RNA, and phospholipids. Most
common in the form of polyprotic phosphoric acid (H3PO4) in soil, but it is taken up most
readily in the form of H2PO4. Phosphorus is limited in most soils because it is released
very slowly from insoluble phosphates. Under most environmental conditions it is the
limiting element because of its small concentration in soil and high demand by plants and
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microorganisms. Plants can increase phosphorus uptake by a mutualism with mycorrhiza.

[4]

A Phosphorus deficiency in plants is characterized by an intense green coloration in

leaves. If the plant is experiencing high phosphorus deficiencies the leaves may become
denatured and show signs of necrosis. Occasionally the leaves may appear purple from an
accumulation of anthocyanin. Because phosphorus is a mobile nutrient, older leaves will
show the first signs of deficiency.
High phosphorus content fertilizers, such as bone meal, is useful to apply to perennials to
help with successful root formation.[4]

Potassium

Potassium regulates the opening and closing of the stomata by a potassium ion pump.
Since stomata are important in water regulation, potassium reduces water loss from the
leaves and increases drought tolerance. Potassium deficiency may cause necrosis or
interveinal chlorosis. K+ is highly mobile and can aid in balancing the anion charges
within the plant. It also has high solubility in water and leaches out of soils that rocky or
sandy that can result in potassium deficiency. It serves as an activator of enzymes used in
photosynthesis and respiration[4] Potassium is used to build cellulose and aids in
photosynthesis by the formation of a chlorophyll precursor.
Potassium deficiency may result in higher risk of pathogens, wilting, chlorosis, brown
spotting, and higher chances of damage from frost and heat.

Nitrogen

Nitrogen is an essential component of all proteins. Nitrogen deficiency most often results
in stunted growth, slow growth, and chlorosis. Nitrogen deficient plants will also exhibit
a purple appearance on the stems, petioles and underside of leaves from an accumulation
of anthocyanin pigements[4]
Most of the nitrogen taken up by plants is from the soil in the forms of NO3–. Amino
acids and proteins can only be built from NH4+ so NO3– must be reduced. Under many
agricultural settings, nitrogen is the limiting nutrient of high growth. Some plants require
more nitrogen than others, such as corn (Zea mays). Because nitrogen is mobile, the older
leaves exhibit chlorosis and necrosis earlier than the younger leaves. Soluble forms of
nitrogen are transported as amines and amides[4]

Sulphur

Sulphur is a structural component of some amino acids and vitamins, and is essential in
the manufacturing of chloroplasts.

Calcium

Calcium regulates transport of other nutrients into the plant and is also involved in the
activation of certain plant enzymes. Calcium deficiency results in stunting.
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Magnesium

Magnesium is an important part of chlorophyll, a critical plant pigment important in

photosynthesis. It is important in the production of ATP through its role as an enzyme
cofactor. There are many other biological roles for magnesium—see Magnesium in
biological systems for more information. Magnesium deficiency can result in interveinal
chlorosis.

Silicon

Silicon is deposited in cell walls and contributes to its mechanical properties including
rigidity and elasticity[citation needed]

[edit] Micronutrients

Iron

Iron is necessary for photosynthesis and is present as an enzyme cofactor in plants. Iron
deficiency can result in interveinal chlorosis and necrosis.

Molybdenum

Molybdenum is a cofactor to enzymes important in building amino acids.

Boron

Boron is important for binding of pectins in the RGII region of the primary cell wall,
secondary roles may be in sugar transport, cell division, and synthesizing certain
enzymes. Boron deficiency causes necrosis in young leaves and stunting.

Copper

Copper is important for photosynthesis. Symptoms for copper deficiency include

chlorosis. Involved in many enzyme processes. Necessary for proper photosythesis.
Involved in the manufacture of lignin (cell walls). Involved in grain production.

Manganese

Manganese is necessary for building the chloroplasts. Manganese deficiency may result
in coloration abnormalities, such as discolored spots on the foliage.

Sodium

Sodium is involved in the regeneration of phosphoenolpyruvate in CAM and C4 plants. It

can also substitute for potassium in some circumstances.
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Zinc

Zinc is required in a large number of enzymes and plays an essential role in DNA
transcription. A typical symptom of zinc deficiency is the stunted growth of leaves,
commonly known as "little leaf" and is caused by the oxidative degradation of the growth
hormone auxin.

Nickel

In higher plants, Nickel is essential for activation of urease, an enzyme involved with
nitrogen metabolism that is required to process urea. Without Nickel, toxic levels of urea
accumulate, leading to the formation of necrotic lesions. In lower plants, Nickel activates
several enzymes involved in a variety of processes, and can substitute for Zinc and Iron
as a cofactor in some enzymes.[2]

Chlorine

Chlorine is necessary for osmosis and ionic balance; it also plays a role in photosynthesis.

Cobalt has proven to be beneficial to at least some plants, but is essential in others, such as
legumes where it is required for nitrogen fixation for the symbiotic relationship it has with
nitrogen-fixing bacteria. Vanadium may be required by some plants, but at very low
concentrations. It may also be substituting for molybdenum. Selenium and sodium may also be
beneficial. Sodium can replace potassium's regulation of stomatal opening and closing[4]

WHAT IS PLANT NUTRITION?

Plants use inorganic minerals for nutrition, whether grown in the field or in a container. Complex
interactions involving weathering of rock minerals, decaying organic matter, animals, and
microbes take place to form inorganic minerals in soil. Roots absorb mineral nutrients as ions in
soil water. Many factors influence nutrient uptake for plants. Ions can be readily available to
roots or could be "tied up" by other elements or the soil itself. Soil too high in pH (alkaline) or
too low (acid) makes minerals unavailable to plants.

FERTILITY OR NUTRITION
The term "fertility" refers to the inherent capacity of a soil to supply nutrients to plants in
adequate amounts and in suitable proportions. The term "nutrition" refers to the interrelated steps
by which a living organism assimilates food and uses it for growth and replacement of tissue.
Previously, plant growth was thought of in terms of soil fertility or how much fertilizer should be
added to increase soil levels of mineral elements. Most fertilizers were formulated to account for
deficiencies of mineral elements in the soil. The use of soilless mixes and increased research in
nutrient cultures and hydroponics as well as advances in plant tissue analysis have led to a
broader understanding of plant nutrition. Plant nutrition is a term that takes into account the
interrelationships of mineral elements in the soil or soilless solution as well as their role in plant
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growth. This interrelationship involves a complex balance of mineral elements essential and
beneficial for optimum plant growth.

ESSENTIAL VERSUS BENEFICIAL

The term essential mineral element (or mineral nutrient) was proposed by Arnon and Stout
(1939). They concluded three criteria must be met for an element to be considered essential.
These criteria are: 1. A plant must be unable to complete its life cycle in the absence of the
mineral element. 2. The function of the element must not be replaceable by another mineral
element. 3. The element must be directly involved in plant metabolism. These criteria are
important guidelines for plant nutrition but exclude beneficial mineral elements. Beneficial
elements are those that can compensate for toxic effects of other elements or may replace
mineral nutrients in some other less specific function such as the maintenance of osmotic
pressure. The omission of beneficial nutrients in commercial production could mean that plants
are not being grown to their optimum genetic potential but are merely produced at a subsistence
level. This discussion of plant nutrition includes both the essential and beneficial mineral
elements.

WHAT ARE THE MINERAL ELEMENTS?

There are actually 20 mineral elements necessary or beneficial for plant growth. Carbon (C),
hydrogen (H), and oxygen (O) are supplied by air and water. The six macronutrients, nitrogen
(N), phosphorus (P), potassium (K), calcium (Ca), magnesium (Mg), and sulfur (S) are required
by plants in large amounts. The rest of the elements are required in trace amounts
(micronutrients). Essential trace elements include boron (B), chlorine (Cl), copper (Cu), iron
(Fe), manganese (Mn), sodium (Na), zinc (Zn), molybdenum (Mo), and nickel (Ni). Beneficial
mineral elements include silicon (Si) and cobalt (Co). The beneficial elements have not been
deemed essential for all plants but may be essential for some. The distinction between beneficial
and essential is often difficult in the case of some trace elements. Cobalt for instance is essential
for nitrogen fixation in legumes. It may also inhibit ethylene formation (Samimy, 1978) and
extend the life of cut roses (Venkatarayappa et al., 1980). Silicon, deposited in cell walls, has
been found to improve heat and drought tolerance and increase resistance to insects and fungal
infections. Silicon, acting as a beneficial element, can help compensate for toxic levels of
manganese, iron, phosphorus and aluminum as well as zinc deficiency. A more holistic approach
to plant nutrition would not be limited to nutrients essential to survival but would include
mineral elements at levels beneficial for optimum growth. With developments in analytical
chemistry and the ability to eliminate contaminants in nutrient cultures, the list of essential
elements may well increase in the future.

THE MINERAL ELEMENTS IN PLANT PRODUCTION

The use of soil for greenhouse production before the 1960s was common. Today a few growers
still use soil in their mixes. The bulk of production is in soilless mixes. Soilless mixes must
provide support, aeration, nutrient and moisture retention just as soils do, but the addition of
fertilizers or nutrients are different. Many soilless mixes have calcium, magnesium, phosphorus,
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sulfur, nitrogen, potassium and some micronutrients incorporated as a pre-plant fertilizer.

Nitrogen and potassium still must be applied to the crop during production. Difficulty in
blending a homogenous mix using pre-plant fertilizers may often result in uneven crops and
possible toxic or deficient levels of nutrients. Soilless mixes that require addition of micro and
macronutrients applied as liquid throughout the growth of the crop, may actually give the grower
more control of his crop. To achieve optimum production, the grower can adjust nutrient levels
to compensate for other environmental factors during the growing season. The absorption of
mineral ions is dependent on a number of factors in addition to weather conditions. These
include the cation exchange capacity or CEC and the pH or relative amount of hydrogen (H+) or
hydroxyl ions (OH-) of the growing medium, and the total alkalinity of the irrigation water.

CEC OR CATION EXCHANGE CAPACITY

The Cation Exchange Capacity refers to the ability of the growing medium to hold exchangeable
mineral elements within its structure. These cations include ammonium nitrogen, potassium,
calcium, magnesium, iron, manganese, zinc and copper. Peat moss and mixes containing bark,
sawdust and other organic materials all have some level of cation exchange capacity.

pH: WHAT DOES IT MEAN?

The term pH refers to the alkalinity or acidity of a growing media water solution. This solution
consists of mineral elements dissolved in ionic form in water. The reaction of this solution
whether it is acid, neutral or alkaline will have a marked effect on the availability of mineral
elements to plant roots. When there is a greater amount of hydrogen H+ ions the solution will be
acid (<7.0). If there is more hydroxyl OH- ions the solution will be alkaline (>7.0). A balance of
hydrogen to hydroxyl ions yields a pH neutral soil (=7.0). The range for most crops is 5.5 to 6.2
or slightly acidic. This creates the greatest average level for availability for all essential plant
nutrients. Extreme fluctuations of higher or lower pH can cause deficiency or toxicity of
nutrients.

THE ELEMENTS OF COMPLETE PLANT NUTRITION

The following is a brief guideline of the role of essential and beneficial mineral nutrients that are
crucial for growth. Eliminate any one of these elements, and plants will display abnormalities of
growth, deficiency symptoms, or may not reproduce normally.

Macronutrients

Nitrogen is a major component of proteins, hormones, chlorophyll, vitamins and enzymes

essential for plant life. Nitrogen metabolism is a major factor in stem and leaf growth (vegetative
growth). Too much can delay flowering and fruiting. Deficiencies can reduce yields, cause
yellowing of the leaves and stunt growth.

Phosphorus is necessary for seed germination, photosynthesis, protein formation and almost all
aspects of growth and metabolism in plants. It is essential for flower and fruit formation. Low pH
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(<4) results in phosphate being chemically locked up in organic soils. Deficiency symptoms are
purple stems and leaves; maturity and growth are retarded. Yields of fruit and flowers are poor.
Premature drop of fruits and flowers may often occur. Phosphorus must be applied close to the
plant's roots in order for the plant to utilize it. Large applications of phosphorus without adequate
levels of zinc can cause a zinc deficiency.

Potassium is necessary for formation of sugars, starches, carbohydrates, protein synthesis and
cell division in roots and other parts of the plant. It helps to adjust water balance, improves stem
rigidity and cold hardiness, enhances flavor and color on fruit and vegetable crops, increases the
oil content of fruits and is important for leafy crops. Deficiencies result in low yields, mottled,
spotted or curled leaves, scorched or burned look to leaves..

Sulfur is a structural component of amino acids, proteins, vitamins and enzymes and is essential
to produce chlorophyll. It imparts flavor to many vegetables. Deficiencies show as light green
leaves. Sulfur is readily lost by leaching from soils and should be applied with a nutrient
formula. Some water supplies may contain Sulfur.

Magnesium is a critical structural component of the chlorophyll molecule and is necessary for
functioning of plant enzymes to produce carbohydrates, sugars and fats. It is used for fruit and
nut formation and essential for germination of seeds. Deficient plants appear chlorotic, show
yellowing between veins of older leaves; leaves may droop. Magnesium is leached by watering
and must be supplied when feeding. It can be applied as a foliar spray to correct deficiencies.

Calcium activates enzymes, is a structural component of cell walls, influences water movement
in cells and is necessary for cell growth and division. Some plants must have calcium to take up
nitrogen and other minerals. Calcium is easily leached. Calcium, once deposited in plant tissue,
is immobile (non-translocatable) so there must be a constant supply for growth. Deficiency
causes stunting of new growth in stems, flowers and roots. Symptoms range from distorted new
growth to black spots on leaves and fruit. Yellow leaf margins may also appear.

Micronutrients

Iron is necessary for many enzyme functions and as a catalyst for the synthesis of chlorophyll. It
is essential for the young growing parts of plants. Deficiencies are pale leaf color of young
leaves followed by yellowing of leaves and large veins. Iron is lost by leaching and is held in the
lower portions of the soil structure. Under conditions of high pH (alkaline) iron is rendered
unavailable to plants. When soils are alkaline, iron may be abundant but unavailable.
Applications of an acid nutrient formula containing iron chelates, held in soluble form, should
correct the problem.

Manganese is involved in enzyme activity for photosynthesis, respiration, and nitrogen

metabolism. Deficiency in young leaves may show a network of green veins on a light green
background similar to an iron deficiency. In the advanced stages the light green parts become
white, and leaves are shed. Brownish, black, or grayish spots may appear next to the veins. In
neutral or alkaline soils plants often show deficiency symptoms. In highly acid soils, manganese
may be available to the extent that it results in toxicity.
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Boron is necessary for cell wall formation, membrane integrity, calcium uptake and may aid in
the translocation of sugars. Boron affects at least 16 functions in plants. These functions include
flowering, pollen germination, fruiting, cell division, water relationships and the movement of
hormones. Boron must be available throughout the life of the plant. It is not translocated and is
easily leached from soils. Deficiencies kill terminal buds leaving a rosette effect on the plant.
Leaves are thick, curled and brittle. Fruits, tubers and roots are discolored, cracked and flecked
with brown spots.

Zinc is a component of enzymes or a functional cofactor of a large number of enzymes including

auxins (plant growth hormones). It is essential to carbohydrate metabolism, protein synthesis and
internodal elongation (stem growth). Deficient plants have mottled leaves with irregular chlorotic
areas. Zinc deficiency leads to iron deficiency causing similar symptoms. Deficiency occurs on
eroded soils and is least available at a pH range of 5.5 - 7.0. Lowering the pH can render zinc
more available to the point of toxicity.

Copper is concentrated in roots of plants and plays a part in nitrogen metabolism. It is a

component of several enzymes and may be part of the enzyme systems that use carbohydrates
and proteins. Deficiencies cause die back of the shoot tips, and terminal leaves develop brown
spots. Copper is bound tightly in organic matter and may be deficient in highly organic soils. It is
not readily lost from soil but may often be unavailable. Too much copper can cause toxicity.

Molybdenum is a structural component of the enzyme that reduces nitrates to ammonia. Without
it, the synthesis of proteins is blocked and plant growth ceases. Root nodule (nitrogen fixing)
bacteria also require it. Seeds may not form completely, and nitrogen deficiency may occur if
plants are lacking molybdenum. Deficiency signs are pale green leaves with rolled or cupped
margins.

Chlorine is involved in osmosis (movement of water or solutes in cells), the ionic balance
necessary for plants to take up mineral elements and in photosynthesis. Deficiency symptoms
include wilting, stubby roots, chlorosis (yellowing) and bronzing. Odors in some plants may be
decreased. Chloride, the ionic form of chlorine used by plants, is usually found in soluble forms
and is lost by leaching. Some plants may show signs of toxicity if levels are too high.

Nickel has just recently won the status as an essential trace element for plants according to the
Agricultural Research Service Plant, Soil and Nutrition Laboratory in Ithaca, NY. It is required
for the enzyme urease to break down urea to liberate the nitrogen into a usable form for plants.
Nickel is required for iron absorption. Seeds need nickel in order to germinate. Plants grown
without additional nickel will gradually reach a deficient level at about the time they mature and
begin reproductive growth. If nickel is deficient plants may fail to produce viable seeds.

Sodium is involved in osmotic (water movement) and ionic balance in plants.

Cobalt is required for nitrogen fixation in legumes and in root nodules of nonlegumes. The
demand for cobalt is much higher for nitrogen fixation than for ammonium nutrition. Deficient
levels could result in nitrogen deficiency symptoms.
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Silicon is found as a component of cell walls. Plants with supplies of soluble silicon produce
stronger, tougher cell walls making them a mechanical barrier to piercing and sucking insects.
This significantly enhances plant heat and drought tolerance. Foliar sprays of silicon have also
shown benefits reducing populations of aphids on field crops. Tests have also found that silicon
can be deposited by the plants at the site of infection by fungus to combat the penetration of the
cell walls by the attacking fungus. Improved leaf erectness, stem strength and prevention or
depression of iron and manganese toxicity have all been noted as effects from silicon. Silicon has
not been determined essential for all plants but may be beneficial for many.

In its purest form, the term "biotechnology" refers to the use of living organisms or their
products to modify human health and the human environment. Prehistoric biotechnologists did
this as they used yeast cells to raise bread dough and to ferment alcoholic beverages, and
bacterial cells to make cheeses and yogurts and as they bred their strong, productive animals to
make even stronger and more productive offspring.

Throughout human history, we have learned a great deal about the

different organisms that our ancestors used so effectively. The
marked increase in our understanding of these organisms and their
cell products gains us the ability to control the many functions of
various cells and organisms. Using the techniques of gene splicing
and recombinant DNA technology, we can now actually combine the
genetic elements of two or more living cells. Functioning lengths of
DNA can be taken from one organism and placed into the cells of
another organism. As a result, for example, we can cause bacterial
cells to produce human molecules. Cows can produce more milk for the same amount of feed.
And we can synthesize therapeutic molecules that have never before existed.
12

The tenure biotechnology refers to the use of systematic techniques, together with genetic
engineering, to urge or cgange plants, animals, and microorganisms. In the most simple forms,
biotechnology has been in use for millennia. For example, Middle Easterners who trained and
bred deer, antelope, and sheep as early as 18,000 B.C.E.; Egyptians who done booze in 4000
B.C.E.; and Louis Pasteur, who grown pasteurization  in 1861, all used biotechnology. In new
years, however, food biotechnology has turn synonymous with the conditions genetically
engineered dishes and genetically mutated mammal (GMO).

Traditional biotechnology uses techniques such as crossbreeding, fermentation, and enzymatic

treatments to furnish preferred changes in plants, animals, and foods. Crossbreeding plants or
animals involves the resourceful thoroughfare of fascinating genes from a single era to another.
Microbial distillation is used in creation booze and alternative alcoholic beverages, yogurt, and
most cheeses and breads. Using enzymes as food additives is an a single more normal form of
biotechnology. For example, papain, an chemical substance performed from papaya fruit, is used
to tenderize beef and explain beverages.
Genetic Engineering

The DNA contained in genes determines hereditary characteristics. Modifying DNA to remove,
add, or change genetic report is called genetic alteration or genetic engineering. In the early
1980s, scientists grown recombinant DNA techniques which authorised them to remove DNA
from a single class and insert it in to another. Refinements in these techniques have authorised
marker of specific genes inside of DNA—and the send of which sold gene method of DNA in to
an a single more species. For example, the genes obliged for producing insulin in humans have
been removed and extrinsic in to bacteria. The insulin which is afterwards constructed by these
bacteria, which is matching to tellurian insulin, is afterwards removed and since to people who
have diabetes. Similarly, the genes which furnish chymosin, an chemical substance which is
concerned in cheese manufacturing, have additionally been extrinsic in to bacteria. Now, instead
of carrying to remove chymosin from the stomachs of cows, it is done by bacteria. This sort of
focus of genetic engineering has not been really controversial. However, applications involving
the use of plants have been some-more controversial.

Among the initial blurb applications of genetically engineered dishes was a chopped tomatoes in
which the gene which produces the chemical substance obliged for softening was incited off. The
chopped tomatoes could afterwards be authorised to rise upon the vine yet removing as well
soothing to be finished and shipped. As of 2002, over forty food crops had been mutated
regulating recombinant DNA technology, together with pesticide-resistant soybeans, virus-
resistant squash, frost-resistant strawberries, corn and potatoes containing a healthy pesticide,
and rice containing beta-carotene. Consumer negativity toward biotechnology is increasing, not
usually in the United States, yet additionally in the United Kingdom, Japan, Germany, and
France, notwithstanding increasing consumer hold of biotechnology. The element objections to
biotechnology and dishes constructed regulating genetic alteration are: regard about probable
mistreat to tellurian illness (such as allergic responses to a “foreign gene”), probable disastrous
stroke to the environment, a ubiquitous confusion about the “unnatural” standing of
biotechnology, and eremite concerns about modification.
Biotechnology in Animals
13

The most argumentative applications of biotechnology engage the use of animals and the send of
genes from animals to plants. The initial animal-based focus of biotechnology was the
capitulation of the use f bacterially Scientists extrinsic daffodil genes and alternative genetic
element in to typical rice to have this golden rice. The outcome is a aria of rice which provides
vitamin A, a nutritious blank from the diets of most people who rely upon rice as a food staple.
[AP/Wide World Photos. Reproduced by permission.] Scientists extrinsic daffodil genes and
alternative genetic element in to typical rice to have this golden rice. The outcome is a aria of
rice which provides vitamin A, a nutritious blank from the diets of most people who rely upon
rice as a food staple. [AP/Wide World Photos. Reproduced by permission.] constructed cow
somatotropin (bST) in dairy cows. Bovine somatotropin, a of course occurring hormone,
increases divert production. This focus has not been commercially successful, however,
essentially since of the expense. The cloning of animals is an a single more intensity focus of
biotechnology. Most experts hold which animal applications of biotechnology will start solemnly
since of the amicable and reliable concerns of consumers.
Concerns about Food Production

Some concerns about the use of biotechnology for food prolongation embody probable allergic
reactions to the eliminated protein. For example, if a gene from Brazil nuts which produces an
allergen were eliminated to soybeans, an particular who is allergic to Brazil nuts competence
right away additionally be allergic to soybeans. As a result, companies in the United States
which rise genetically engineered dishes contingency denote to the U.S. Food and Drug
Administration (FDA) which they did not send proteins which could outcome in food allergies.
When, in fact, a association attempted to send a gene from Brazil nuts to soybeans, the
company’s tests suggested which they had eliminated a gene for an allergen, and work upon the
plan was halted. In 2000 a code of taco shells was detected to enclose a accumulation of
genetically engineered corn which had been authorized by the FDA for use in animal feed, yet
not for tellurian consumption. Although multiform antibiotechnology groups used this situation
as an e.g. of intensity allergenicity stemming from the use of biotechnology, in this box the
protein constructed by the genetically mutated gene was not an allergen. This situation
additionally demonstrated the difficulties in gripping lane of a genetically mutated food which
looks matching to the unmodified food. Other concerns about the use of recombinant DNA
record embody intensity waste of biodiversity and disastrous impacts upon alternative aspects of
the environment.
Safety and Labeling

In the United States, the FDA has ruled which dishes constructed yet biotechnology need the
same capitulation routine as all alternative food, and which there is no fundamental illness risk in
the use of biotechnology to rise plant food products. Therefore, no tag is compulsory simply to
brand dishes as products of biotechnology. Manufacturers bear the weight of explanation for the
reserve of the food. To support them with this, the FDA grown a decision-tree proceed which
allows food processors to expect reserve concerns and know when to deliberate the FDA for
guidance. The preference tree focuses upon toxicants which have been evil of any class involved;
the intensity for transferring food allergens from a single food source to another; the
thoroughness and bioavailability of nutrients in the food; and the reserve and nutritive worth of
newly introduced proteins.
14

Labeling of genetically mutated dishes has sparked a single more debate. Labels have been
compulsory upon food constructed by biotechnology to surprise consumers of any intensity
illness or reserve risk. For example, a tag is compulsory if a intensity allergen is introduced in to
a food product. A tag is additionally compulsory if a food is remade so which the nutritious calm
no longer resembles the strange food. For example, supposed golden rice has been genetically
engineered to have a aloft thoroughness of beta-carotene than unchanging rice, and to illustrate it
contingency be enclosed upon the label. In reply to consumer demands, regulators in England
have instituted imperative labeling laws for all finished dishes and menus containing genetically
mutated ingredients. Similar yet reduction limiting laws have been instituted in Japan. In Canada,
the process upon labeling has remained identical to which of the United States.

Some consumer advocates say which not requiring a tag upon all genetically mutated dishes
violates consumers’ right to have sensitive food choices, and most producers of sure foods, such
as dishes containing soy protein, right away embody the tenure “non-GMO” upon the tag to
prove which the product does not enclose genetically mutated ingredients.

The focus of recombinant DNA record to foods, ordinarily called biotechnology, might be
noticed as an prolongation of normal cross-breeding and distillation techniques. The record
enables scientists to send genetic element from a single class to another, and might furnish food
crops and animals which have been opposite than those performed regulating normal techniques.
The FDA has determined procedures for capitulation of food products made regulating
recombinant DNA record which need food producers to denote the reserve of their products. The
American Dietetic Association, the American Medical Association, and the World Health
Organization have any adopted statements which techniques of biotechnology might have the
intensity to urge the food supply. These organizations and others admit which long-term illness
and environmental impacts of the record have been not known, and they inspire redundant
monitoring of intensity impacts.\

Introduction

The origin of the flowering plants was described by Charles Darwin as "that abominable
mystery". Flowering plants are the dominant plants on the earth today with an estimated quarter
of a million species. Unlike lower plants like ferns and seaweeds which rely on spores to invade
new habitats, flowering plants are dispersed by seeds and this undoubtedly accounts for their
success. Lower plants alternate in their life cycle between distinct sporophyte and gametophyte
stages. While these phases can be recognised in flowering plants as well, an understanding of
this is not required for CAPE Biology and these concepts are best ignored at this level.
 

Flower structure

The flower is simply a collection of highly modified leaves. The outermost series is
collectively termed the perianth and comprises sepals, which protect the developing
flower, and petals, which often serve to attract pollinators. Within the perianth lie the
15

actual reproductive organs. The stamens produce pollen from which the male gametes
are ultimately derived. The carpels contain an ovary with ovules in which the female
gametes develop. 

Following the transfer of  pollen to the stigma or receptive surface of the carpel, in  a
process called pollination, nuclei  from the pollen will ultimately fuse with nuclei
within the ovule's embryosac to form the seed.

In some flowering plants

all the perianth parts are
similar; there are no
separate sepals and petals.
This can often be seen in
lily-like plants like the
Spider Lily (Hymenocallis
caribaea).

The flowering plants show great variation in flower structure and these characteristics are central
to flowering plant classification. In most cases, flowers are not borne singly but grouped on a
common stalk and termed an inflorescence. Various inflorescence types are recognised and
while these do not concern us at this level be aware that these are very important in
classification.
 
 
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Sometimes the flower

has closely associated
with it a prominent
(often coloured) leaf
termed a bract. In
bougainvillea (left)
these bracts are more
attractive than the
actual flower which is
pale and
inconspicuous.
Similarly, red bracts
constitute the most
conspicuous part of the
inflorescence of the red
ginger lily (right). 

Overview of sexual reproduction

The details of pollen grain development and embryosac formation will not be discussed here as
these are well covered in all textbooks. Some points, however, are worth emphasising.

 In both pollen and ovule development, meiosis occurs which not only halves the
chromosome complement of cells but involves crossing over and so a reshuffling of
genes, ensuring genetic  recombination.

 Unlike other seed plants, the ovules are completely sealed within the carpel. This means
that the pollen tube must actually bore its way down through the style into the ovary and
this has led to the evolution in flowering plants of self incompatibility mechanisms to
prevent the entry of "undesirable" pollen.

 The flowering plants are unique in having a double fertilisation event. One nucleus from
the pollen grain will fuse with the egg to form the diploid zygote which develops into a
minature plant, the embryo. More unusual is the fact that a second nucleus from the
pollen will fuse with 2 polar nuclei of the embryosac to form a triploid cell which
develops into the endosperm, a tissue that will provide nutrition for the embryo. The
seed that results then comprises the embryo, a minature plant ready to start a life of its
own, with reserve materials either in a surrounding endosperm or transferred to the
embryo's cotyledons, all within a tough seed coat.
17

 One final unique feature of sexual reproduction in flowering plants is the development of
fruit. Following fertilisation, as each ovule form a seed, the ovary swells to form the
fruit. The biologist's definition of a fruit is simply an organ for dispersing seeds. This is
quite different to the layman's concept of a fruit as something soft, juicy and edible. Fruit
may split or remain intact. They can be hard and dry like a calabash or fleshy like a
tomato or a cucumber. There is a complex system of recognised fruit types but you need
not worry about these in this course.

Promotion of cross-fertilisation

To ensure the long term survival of any species it is advantageous to have a large and varied
population on which natural selection can act. Such variability is promoted by cross-fertilisation
and reduced by self-fertilisation. Not surprisingly flowering plants have evolved a number of
mechanisms which maximise cross-pollination.

1. Separation of the sex organs

Some plants have unisexual flowers (i.e. contain either carpels or stamens but not both) rather
than bisexual flowers. In dioecious species, plants are either male or female (e.g. nutmeg,
marijuana). In monoecious species, there are separate male and female flowers but on the same
plant (e.g. castor oil, breadfruit, corn, pumpkin).

Female & male  flowers of Jatropha

integerrima, a common garden shrub in the Castor oil, Ricinus communis
Caribbean

 
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2. Timing mechanisms
Sex organs may mature at different
times. Protandry is where the anthers
mature first while protogyny is where
the stigma is receptive before the anthers
mature. Both avocado and soursop are
protogynous. The Anthurium bloom,
which is not a flower but an
inflorescence or collection of flowers, is
another example. The fleshy spadix feels
wet when the minute female flowers are
ripe and dry and powdery when the
pollen is being shed. Male and female
parts are never ripe together. 

Soursop flower with perianth (left),

without perianth (centre), with anthers
shed (right)

3. Specialised pollination mechanisms

Many plants have highly specialised pollination
mechanisms, often adapted to a single insect, bird or bat
pollinator. The highly specialised flower structure and
manipulation of the behaviour of the vector ensure cross-
pollination. Click here to see flowers with insect eyes! 

Aristolochia, Dutchman's pipe,

is a woody vine with several
species native to the
Caribbean, all with strange,
bent, tubular flowers. Flies are
attracted by a putrid scent
emitted by the flower and
enter its tunnel-like interior.
Downward pointing hairs
allow them to proceed further
but not escape. The flowers
are protogynous so on this first
day the stigmas are receptive
but no pollen is shed. On Day
2, the stigmas wither and the
pollen is now shed, coating the
Several Aristolochia flower types entrapped flies. The next day
the hairs blocking the flies'
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escape whither, setting the

"prisoners" free. They now
visit a new freshly-open,
"fragrant" flower. The story is
repeated again but on arrival
they bring pollen to the 
receptive stigmas of these Day
1 flowers and so achieve
cross-pollination! 

Orchids also provide good

examples of highly evolved
flower-pollinator relationships
- the stuff of natural history
documentaries.

4. Heterostyly 
In some species a plant bears one of two
morphological forms of a flower.
Flowers may have long styles with
anthers near the flower base (pin form)
or short styles with anthers borne on
high (thrum form). These arrangements
prevent selfing when a pollinator visits
and ensures cross-pollination between
plants with the different flower types.
The Red Cordia, Cordia sebestena,
(right) shows this.

 
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5. Self incompatibility 
When a pollen grain germinates on a stigma of the same species a
recognition process takes place. This involves molecules from the
stigma entering the pollen grain and vice versa. This process is
under genetic control and results in pollen from the same plant or
a different plant of similar genetic make-up being rejected. The
pollen grain may be prevented from germinating or the growth of
its tube may be inhibited within the style. Many plants including
cabbage and tobacco show this. 
 

Asexual reproduction

Sexual reproduction in flowering plants involves meiosis and the fusion of gametes, usually from
different plants, and results in seedlings which differ from their parents. In asexual reproduction,
exact copies of the parent plant result as the new plants are formed simply by mitosis. Sometimes
flowers are involved in this process but usually only the vegetative parts of the plant are involved
and so this is termed vegetative reproduction.
 
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Plants have a
range of
perennating
organs, which
store reserves and
are produced to
enable the plant to
tide over periods
of unfavourable
environmental
conditions, e.g. the
dry season in the
tropics, winter in
temperate regions.
Examples of these
are the rhizomes
(underground
stems) of ginger,
the stem tubers of
English potato, the
corms (vertical
stem tubers) of
eddo, the root
tubers of yam and
sweet potato and
the bulbs of
onions and lily-
like plants. These
can be used as a
means of 
propagating the
plant in question.

corm bulb

Plants may also send out creeping branches or runners (Water lettuce), or may produce
adventitious buds along the edges of leaves (Bryophyllum sp.). Most Agave spp. (Maypole,
Dagger, Lapitte) have sterile flowers but then produce minature plants or bulbils on the pole-like
inflorescence.
 
 
22

Water lettuce, Pistia stratioites Bryophyllum sp. Agave sp.

Propagation

 
Artificial vegetative reproduction at its simplest involves inducing
stem cuttings to form roots by maintaining these in a humid
environment. Treating the cuttings with a synthetic auxin such as
indole-3-butyric acid (typically diluted with inert talcum powder)
often speeds up the induction of adventitious roots. If cuttings do
not readily root it may be necessary to use the technique of air-
layering where rooting is induced on a branch still attached to the
parent plant. This involves removing a ring of bark, packing the
exposed area with moist soil and keeping this sealed with plastic
film until roots form, at which time the rooted cutting is removed
and planted. Some woody cuttings are very difficult to root. In
such cases, one must graft  a bud (budding) or shoot cutting
(grafting) onto a root stock. Such vegetative propagation is
necessary when an exact copy of the plant is desired. This is true
for particular varieties (more correctly cultivars) of woody plants
like  mango and hibiscus. Since sexual reproduction leads to
recombination, seed from such plants will not produce offspring
with identical traits to the parent and vegetative propagation is
required. 
 
23

In plant tissue culture, sterile

portions (explants) of a plant are
transferred to sterile medium
containing agar, mineral
nutrients, vitamins, sucrose and
plant hormones like auxins and
cytokinins. This induces cells to
return to a meristematic dividing
phase and produce a disorganised
proliferation of cells termed a
callus. By later changing the
culture medium, calli can then be
induced to form shoots and roots
and so regenerate lots of tiny
plants. This technique can
propagate plants on an industrial
scale and has been used in the
Caribbean for banana
propagation. It does require
skilled personnel, specialised
facilities and can take a
considerable time to regenerate
plantlets and then to grow them to
a  field-ready stage. It is also
well-documented that genetic
variation, termed somaclonal
variation, can develop during the
tissue culture process so that the
final plants may not be identical.
These techniques of in vitro
propagation are especially
important for the generation of
disease-free planting material.
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