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BIOCHEMICAL CHANGES OF
CARBOHYDRATES IN FOOD
DR. FESTUS ACQUAH

OUTLINE/OBJECTIVES
At the end of this topic, the student should be able to understand and make
practical applications in the following areas:

• Structure of Carbohydrates….Understand the chemical structure of


Carbohydrates.

• Metabolism of Complex Carbohydrates.

• Changes in Carbohydrates during seed germination

• Application in food processing

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CARBOHYDRATES
• One of the most abundant class of biomolecules on earth.
• produced by the photosynthetic activity of the green plants
• also referred to as saccharides because of the sweet taste of many
carbohydrates (Latin, saccharum, meaning sugar)
• They were once thought to be “hydrates of carbon” hence the name
carbohydrates.

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CARBOHYDRATE SOURCES
• Cereals – Wheat, Maize, Rice, Barley, Sorgum
• Sugar Crops – Sugar Cane, Sugar beet
• Root Crops – Sweet Potatoes
• Pulses – Soya beans, Kidney beans
• Fruits - Banana
• Vegetables - Carrots
• Milk Products

CARBOHYDRATES
WHAT ARE THEY?
• Carbohydrates are polyhydroxy aldehydes or polyhydroxy
ketones or compounds that produce such substances upon
hydrolysis.
• So they are organic compounds composed of Carbon, Hydrogen
and oxygen.
• They also contain nitrogen, sulphur and phosphorus.
• general Formula: CnH2nOn or Cn(H2O)n

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CARBOHYDRATES
• Polyhydroxyl: Having more than one hydroxyl group.
(-OH)
H-C=O
• Aldehyde Group (H-C=O) 
H-C-OH

=o

OH-C-H
• Keto Group (C-C-C) 
H-C-OH

CH2OH
CH2OH

C=O

OH-C-H

H-C-OH

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Classification of carbohydrates
• Carbohydrates are mainly classified into four different groups:
• Monosaccharides
• Disaccharides
• Oligosaccharides
• Polysaccharides

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MONOSACCHARIDES

• Monosaccharides are those carbohydrates which Cannot


be Hydrolyzed further into more simple carbohydrates.
• Thus, they are the Simplest form of Carbohydrates.
• simple sugars, consist of a single polyhydroxy aldehyde or
ketone unit.
• Familiar examples are:
Glucose, Fructose, Ribose & Galactose.

Monosaccharides are further classified


on the basis of:
• Carbonyl group (Aldehyde or Ketone) Group:
• Aldomonosaccharides (Aldoses).
• Ketomonosaccharides (Ketoses).
• Carbon Chain Length.
• Trioses.
• Tetroses.
• Pentoses.
• Hexoses.
• Heptoses.

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CARBONYL GROUP
H-C=O CH2OH
 
H-C-OH C=O
 
OH-C-H GLUCOSE OH-C-H
 
H-C-OH FRUCTOSE--> H-C-OH
 
H-C-OH H-C-OH
 
CH2OH CH2OH

CARBON CHAIN
H-C=O
 H-C=O
H-C-OH 
 OH-C-H
OH-C-H 
GLUCOSE H-C-OH

H-C-OH ARABINOSE--> 
 H-C-OH
H-C-OH 
 CH2OH
CH2OH

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EXAMPLES OF MONOSACCHARIDES
NO. OF CARBON ATOMS ALDO KETO

3C TRIOSE GLYCERALDEHYDE DIHYDROXYACETONE

4C TETROSE ERYTHROSE ERYTHRULOSE

5C PENTOSE RIBOSE, XYLOSE RIBULOSE, XYLULLOSE

6C HEXOSE GLUCOSE, GALACTOSE, FRUCTOSE


MANNOSE

Structures of the D-Aldoses

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Structures of the D-Ketoses

Classes of carbohydrates
• Monosaccharides:
• ketomonosaccharides of more than four carbons and aldomonosaccharides
of at least 4 carbons tend to have cyclic structures.

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Classes of carbohydrates
• Disaccharides:
• Made from two monosaccharide units linked together
• The covalent bonds that hold monosaccharide units together are called
glycosidic bonds.

GLYCOSIDIC BONDS
• Are the covalent bonds that link monomers in disaccharides or
polymers of carbohydrates.

• Can be designate alpha (α) or beta (β)depending on the orientation of


the Carbon number 1 (next to the carbonyl carbon in a straight chain
conformation) forming the bond.

• The linkages can be 1-2, 1-3, 1-4, 1-6. In starch, α 1-4 linkage form
straight chains and α 1-6 linkages form branches in the straight chain.

• These linkages can be broken by specific enzymes.

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Sugars - (This refers to the all the monosacharides


and the dissacharides
• PHYSICAL PROPERTIES
• Sweet when dissolved
• Different sugars have different sweetness

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CHEMICAL REACTIONS SUGARS UNDERGO

CHEMICAL REACTIONS SUGARS UNDERGO

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Classes of carbohydrates
• Oligosacharides:
• Made from few (3-10) monosaccharide units linked together by glycosidic bonds and
they may be branched or linear.

Polysaccharides (glycans)
• They have the general formula (C6H10O5)n where ‘n’ is a large number
• sugar polymers containing more than 20 or so monosaccharide units, and
some have hundreds or thousands of units.
• Homopolysaccharides: composed of the same kind of monomer unit eg.
Glucosans (Starch, glycogen, cellulose, dextrin, dextran) and Fructosans
(Inulin).

• Heteropolysaccharides: contain two or more different kind of monomers.


Eg. Hyaluronic Acid of animal and humans (repeating disaccharide units of
glucuronic acid and Nacetylglucosamine), Agar and pectin from plant sources

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DEGREE OF POLYMERISATION (DP)


• DP – Generally Refers to the average number of monomer units per
molecule.
• For Carbohydrates: Refers to the average number of anhydroglucose
units per molecule.

GLYCOSIDIC BONDS
• Are the covalent bonds that link monomers in disaccharides or
polymers of carbohydrates.

• Can be designate alpha (α) or beta (β)depending on the orientation of


the Carbon number 1 (next to the carbonyl carbon in a straight chain
conformation) forming the bond.

• The linkages can be 1-2, 1-3, 1-4, 1-6. In starch, α 1-4 linkage form
straight chains and α 1-6 linkages form branches in the straight chain.

• These linkages can be broken by specific enzymes.

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Examples of Polysaccharides

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FIBRE
Dietary fiber is defined as plant polysaccharides that are indigestible by humans, plus lignin. The
major components of dietary fiber are cellulose, hemicellulose, pectin, hydrocolloids and lignin.

Fibre is present in whole grains, fruits and vegetables, especially the outer covering of seeds.

There are two types of fibre:

• Soluble fibre - found in fruit, vegetables, pulses and oats.

• Insoluble fibre - found in cereals such as bread and pasta.

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FUNCTIONAL PROPERTIES IN FOOD PRODUCTS


• help to cause the colour change of bread, toast and bakery products;

• contribute to the chewiness, colour and sweet flavour of caramel;

• thicken products such as sauces and custards.


• Provide Bulk
• Consistency
• Texture

CHEMICAL REACTIONS OF CARBOHYDRATES IN FOODS


• Caramelization
• Caramelization is a nonenzymatic browning reaction formed by heating carbohydrates like
sucrose or reducing sugars.
• Reactions are facilitated by small quantity of acids, base and certain salts.
• Caramelization is an entirely different process from Maillard browning, though the results of the
two processes are sometimes similar to the naked eye (and taste buds).

• The final product caramel contains a complex mixture of polymeric compound, formed from
unsaturated cyclic compounds.

• Flavour and aroma compounds are also formed.

• Heating causes the dehydration of sugar molecule with introduction of double bonds or
formation of anhydro rings. Intermediates such as 3-deoxy osones and furans are formed.

• The unsaturated rings may condense to form useful, conjugated double-bond containing, brown
coloured polymers.

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Caramelization
• The types of reactions that occur during the
process of caramelization are
• Sucrose inversion to fructose and glucose
• Condensation reactions
• Intramolecular bonding
• Isomerization of aldoses to ketoses
• Dehydration reactions
• Fragmentation reactions
• Unsaturated polymer formation.

CARAMELIZATION
• To make caramel a carbohydrate is heated alone or in the presence of acid,
a base or salt.

• The carbohydrates most often used are sucrose, but fructose, glucose,
invert sugar, malt syrups and molasses may also be used.

• Acid used are food grade sulfuric, sulfurous, phosphoric, acetic and citric
acids.

• Bases that may be used are ammonium, sodium, potassium and calcium
hydroxides.

• Salts that may be used are ammonium, sodium, potassium carbonates,


bicarbonates, phosphates, sulphates or bisulphates.

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CLASSES OF CARAMEL
Class I caramel :Prepared by heating a plain carbohydrate
Class II caramel :Prepared by heating a carbohydrate in the presence of a
sulphite
Class III caramel :Prepared by heating a carbohydrate in the presence of a
source of ammonium ion.
Class IV caramel :Prepared by heating a carbohydrate in the presence of a
both sulphite and ammonium ions

• Caramelization may sometimes cause browning in the same foods in which


the Maillard reaction occurs, but the two processes are distinct.

• They both are promoted by heating, but the Maillard reaction involves
amino acids, while caramelization is simply the pyrolysis of certain sugars.

THE MAILLARD REACTION


• The Maillard reaction is one of the most common and most complex
reaction that takes place mainly in foods, first described by the French
chemist Louis-Camille Maillard in 1912.

• Maillard reaction is the sequence of events that begins with reaction


of the amino group of amino acids with a glycosidic hydroxyl group of
sugars; the sequence terminates with the formation of brown
nitrogenous polymers called melanoidins.

• It is also a non-enzymatic browning process.

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THE MAILLARD REACTION


• The chemical reactions which
occur can be broadly divided
into three main stages:
• 1. Early stage: The carbonyl
group of the sugar reacts with
the amino group of the amino
acid, producing N-substituted
glycosylamine and water. This
is followed by unstable
glycosylamine undergoing
Amadori rearrangement,
forming ketosamines. The
products formed at this stage
are colourless

THE MAILLARD REACTION


• 2. The advanced or intermediate stage:
Degradation of the rearrangement product
via enolization (C1-C2, C2-C3, etc.), and
subsequent secondary reactions. In general,
under acidic conditions, the nitrogen is
protonated, and 1,2-enolization is assisted by
the positively charged nitrogen acting as an
electron sink. On the other hand, alkali and
strong basic amine favour 2,3-enolization.
The reactions of this stage lead to sugar
dehydration, sugar fragmentation and amino
acid degredation. The products of this phase
are colourless or yellow

• 3. The final stage: The reactions of this


stage include aldol condensations,
aldehyde-amine condensations,
formation of heterocyclic compounds
which are the of browned, aroma and
flavoured compounds, and polymers/co-
polymers or melanoidins.

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Other carbohydrate modifications in food


• Dextrinisation
• Dextrinisation occurs when the heat breaks the large starch
polysaccharides into smaller molecules known as dextrins.
Dextrins are typically brown in colour and have a distinct
taste and consistency.
• It is usually brought about by dry roasting the starch either
alone, making use of its natural 10–20% moisture content,
or (industrially) in the presence of catalytic quantities of
acid. This gives rise to a range of polymer fractions of
varying chain length (low conversion).
• The second aspect involves a recombination of these
fragments (repolymerisation) but this time in a branched
manner (high conversion).
• Basically, dextrin can be produced using three different
methods such as enzymatic hydrolysis, acid hydrolysis and
by the action of heat, or both heat and acid, on starch.

Dextrinisation
• High grade dextrins are prepared by heating starch which has been moistened with
small quantity of dilute nitric acid and dried ,at 110-115 OC, the product is known as
white dextrin.
• Inferior dextrins , which have yellow or brown colour , are prepared by roasting
starch at 150-250OC without addition of acid.

• Uses
• 1. As an excipient for dry extracts and pills.
• 2. It is used for preparing emulsions and dry bandages.
• 3. It is employed for thickening of dye-pastes and mordants used in printing fabrics.
• 4. It is used for sizing paper and fabrics.
• 5. It is employed for preparing felt and printing tapestries.
• 6. It is used for preparing printer’s inks, glues and mucilage.
• 7. It is employed for polishing cereals.
• 8. It is extensively used in making matches, fireworks and explosives.

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Other carbohydrate modifications in food


• Gelatinisation
• Gelatinisation When starch is mixed with water and heated (boiled), the starch granules absorb water,
swell and eventually rupture, absorbing liquid to produce a thickened mixture. Starch gelatinization is
the disruption of molecular orderliness within the starch granule. It results in
• granular swelling,
• crystallite melting,
• loss of birefringence,
• viscosity development, and
• Solubilization
• When starch granules are viewed under a microscope with a polarised light, the intact crystalline
structure refracts the light in two different directions producing a “Maltose Cross”. This phenomenon is
called Birefrigence

• The disappearance of these crosses on heating starch can be used to determine the gelatinization
temperature (the temperature at which a particular starch gelatinizes to form a starch paste).

• Starch Paste refers to the viscous colloidal mass resulting from the gelatinization of starch in water.

Starch Gelatinization

C E
D
A

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Starch Gelatinization
• Peak viscosity is reached during the gelatinzation when all
the granuels are swollen to their maximum level.
• Rupturing of the swollen granules at full gelatinization leads
to a reduction in the viscosity below peak viscosity.
• After full gelatinisation and cooling to a particular
temperature, the viscosity of the starch increases again to a
viscosity level called the endpoint viscosity and this process
is called “setback” or retrogradation. This differs for starch
from various starches
• During retrogradation, the amylose or amylopectin
polymers begin to reassociate via hydrogen bonding leading
to the formation of ordered structures which if continues
can lead to more ordered regions.
• Starch with hy amylose content have a much higher
tendency to retrograde.
• Retrogradation results in the expulsion of water (syneresis)
• Amylopectin has a lower tendency to retrograde

Digestion , absorption anD metabolism of carbohyDrate

• The goal is to break sugar and starches into small molecules-chiefly


glucose-that the body can absorb and use.
• A large starch molecule require extensive breakdown.
The disaccharide needs only to be broken once.
And the monosaccharide not at all.
• The initial splitting begins in the mouth, and the final splitting and
absorption occur in small intestine, and the conversion to common
energy (glucose) takes place in the liver and other tissues of the body.

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Digestion
In the mouth:
Mastication: the process by which food is
crushed and ground by teeth .The chewing of
high-fiber food slows eating and stimulate the
flow of saliva.
Starch: salivary glands secretes
saliva into the mouth to moisten the food.
The salivary enzyme amylase begins
digestion.

Starch Amylase small polyshcarrides,


maltose(disacharride)

Fibers: the mechanical action of the


mouth crushes and tears fiber in food
and mixes it with saliva to moisten
it for swallowing.

In the stomach:
Peristalsis: wave like muscular contraction.
The swallowed bolus(a portion of food swallowed
at one time) mixes with stomach acids and protein
digesting enzymes, which inactivate salivary amylase.
Starch: stomach acid inactivates salivary enzymes,
halting starch digestion.
To small extent ,the stomach acid continue breaking
the starch, but it juices contain no enzymes to digest
CHO.
Fibers: is not digested in stomach and delays gastric
emptying thereby provide feeling of fullness and
satiety.

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Small intestines
The final step takes place in the outer membrane of the
intestinal cells. The disaccharide digestion begins at this
point. There specific enzymes secreted from the intestinal
glands breaks down specific disaccharides.

The disaccharide enzymes on the surface of the small


intestinal cells hydrolyze the disaccharides into
monosaccharaides
Maltose maltase Glucose +Glucose
Sucrose sucrase Fructose+Glucose
Lactose lactase Galactose+Glucose
==>intestinal cells absorb these monosaccharaides.

Fibers: is not digested ,and delays the absorption of


other nutrients.

Large intestine:
• Within 1-4 hours after a meal, all of the sugars and most of the starches
have been digested.
• Only fibers remain in the digestive tract.
• Fibers in large I. attracts water, which softens the stool for passage
without straining.
• Also, bacteria in the GI tract ferment some fibers.
• Most fiber passes intact through digestive tract to the large intestine.
Here, bacterial enzyme digest fiber.

Some fibers Bacterial Enzymes short-chain


fatty acids, gas & water

• Colon uses these small fat molecules for energy. Metabolism of short
chain fatty acids occurs in the cells of liver. fiber therefore can contribute
some energy. Depending on the extent to which they are broken down
by bacteria an d the fatty acids are absorbed.

• Fiber holds water ,regulate bowel activity, and bind substances such as
bile ,cholesterol, and some minerals, carrying them out of the body.

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Carbohydrate absorption
• Peristaltic movement moves the monosaccharaides into the jejunum where
digestion is completed and absorption begins.

• Absorption is increases as a result of the intestinal villi(small mucus projections


lining the small intestine). Each villi contain blood capillary into which the
monosaccharaides passes via diffusion or active transport.

• Glucose is unique that it can be absorbed to some extent through the lining of
mouth, but for most part nutrient absorption takes place in the small intestine.

• Glucose and galactose traverse the cell lining the small I. by active transport.

Carbohydrate absorption
• Fructose is absorbed by facilitated diffusion, which slows its entry and produces a
smaller rise in blood glucose.

• Likewise, unbranced chains of starch are digested slowly and produce a smaller
rise in blood glucose than branched chains, which have many more places for
enzymes to attack and release glucose rapidly.

o As the blood from the intestine circulates through the liver, cells there take up
fructose and galactose and convert them to other compounds ,most often to
glucose .

o Thus all disaccharides provide at least one glucose molecule directly ,and they can
provide another one indirectly –through the conversion of fructose and galactose
to glucose.

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Carbohydrate Metabolism
• Metabolism is the all the chemical processes that a cell/organism
undertakes to stay alive.
• It can be a degredative reaction (catabolic) or building up reaction
(anabolic).
• A product of metabolism is called a metabolite
• Glucose plays a central role in metabolism of carbohydrates.
• When glucose gets to the liver, much of it is converted to glycogen for
storage in the liver (Glycogenesis)
• The rest is released into the blood stream to be carried to the other
tissues of the body to supply energy.
• Muscles can also store glucose as glycogen but this is only to supply
the muscle only with energy later.

Carbohydrate Metabolism
• Glucose is the preferred energy source for the brain, other nerve cells etc.
• Some glucose can be broken down to ribose which is used in making
nucleic acids (DNA and RNA)
• Excess glucose can be converted to fat for storage in the adipose tissue.
• After a while (about 4 hours) after a meal, stored glycogen in the liver is
broken down to produce glucose for the body (glycogenolysis).
• When carbohydrate stores are depleted, the body breaks down fat stores
for a little portion of it to be used to produce glucose (gluconeogenesis)
while the liver uses the rest to make ketone bodies, which can be
metabolised by tissues of the body for energy.
• Tissues that can directly use fatty acids for energy also use some of the
fatty acids for energy.

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Carbohydrate Metabolism
• Long periods of not taking carbohydrates can also lead to body
proteins being broken down to form glucose which is needed at all
cost to serve as fuel for some tissues such as the brain that needs
them.
• This synthesis of glucose from non-carbohydrate sources is called
gluconeogenesis

SOME APPLICATIONS OF CARBOHYDRATE


METABOLISM
• Production of egg powder:
• Whole egg powder is the dried form of whole eggs. It is
commonly used as a whole egg substitute in protein-rich
baked goods due to its longer shelf life compared to fresh
eggs.
• Eggs contain approximately 1.0% carbohydrate and it is mainly
glucose.
• Glucose affects adversely the storage stability of dried eggs
and causes undesirable changes in physico-chemical and
functional properties of the powder. Therefore, microbial or
enzymic techniques are used to desugar eggs before drying.
• During the production of egg powder, glucose is removed
from the egg by the addition of the glucose removal enzyme
glucose oxidase (or microbes to ferment the glucose).
• Glucose oxidase oxidizes theβ-d-glucose to pro-duce d-
glucono-δ-lactone and H2O2.
• With glucose removed the chemical reactions such as the
maillard reaction cannot occur in the product.

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METABOLISM IN A GERMINATING SEED

PARTS OF A SEED

Cotyledons store food

(Cotyledon)

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PARTS OF A SEED
1. Inside of the seed coat, an embryo is protected.
2. The embryo is an immature plant with all of the parts of an adult plant.
• It has leaves and roots although this are very small
• The shoot of the embryo are called plumules . Plumules of mononocots
have a protective sheath called the coleoptile.
• The embryonic leaves which becomes the first leaves at germination is
the cotyledon. In dicots it stores the food used during germination.
• The embryonic roots are called radicles. Radicles of mononocots have a
protective sheath called the coleorhiza.
• The embryonic stem is called the hypocotyl
3. The seed is filled with nourishment for the new plant; in monocots
nourishment is stored in the endosperm while in dicots it is stored in the
cotyledons.

SEED DORMANCY
 Factors that affect dormancy
 A waterproof seed coat
 Light
 Temperature
 Hard Seed Coat
 Period after ripening
 Germination inhibitors. Eg. salicylic acid
 Immaturity of the seed embryo
 Impermeability of seed coat to water
 Impermeability of seed coat to oxygen
 Mechanically resistant seed coat
 Presence of high concentrate solutes
 For germination to occur, these must be negated
 Coat broken down
 oxygen available
 Water absorbed
 Growth promoters activated, inhibitors are inhibited.

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• In general, germination process can be


distinguished into three phases:
• phase I: water imbibition by seed;

• phase II: reactivation of metabolism; and

• phase III: radicle protrusion

• To sustain a good seedling development, seed


stores a food reserve mainly as proteins, lipids
and carbohydrates.

Phase 1: water imbibition


 includes imbibition of dry seeds and the early plateau phase of
water uptake.
 Water is the most important factor in germination; an adequate
continuous supply of water is necessary for complete emergence.
 Water functions as
 triggering production of phytohormones (mainly gibberellic acid (GA)) and
enzymes needed for fuel mobilization.
 turgor pressure for moving the radicle root down and the cotelydons up
 and for transporting nutrients and enzymes within the seed

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phase II: reactivation of metabolism and nutrient mobilisation


• On seed hydration, separate intercellular bodies of seed stored
carbohydrates, proteins, lipid and phosphate act as energy
source and carbon skeleton.
• Seed imbibition triggers many metabolic processes such as
activation or freshly synthesis of hydrolytic enzymes which result
in hydrolysis of stored starch, lipid, protein hemicellulose,
polyphosphates and other storage materials into simple available
form for embryo uptake.

• CARBOHYDRATES:
• In the endosperm, α-amylase, β-amylase, debranching enzyme, and α-
glucosidase combine to digest starch. α-Amylase plays the main role in
digesting storage starch, followed by α-glucosidase. β-Amylase and
debranching enzyme are not important at the beginning of germination,
but contribute in a later stage of germination. The main end product
from degradation of starch is maltose. α-Glucosidase in the endosperm
can convert maltose into glucose. These sugars from degradation of
starch are absorbed into the scutellum, where they are converted into
sucrose and transported to the embryonic axis

phase II: reactivation of metabolism and nutrient mobilisation


• Proteins:
• The major storage proteins present in the endosperm are
hydrolyzed by proteinases during germination. Proteins or
polypeptides are hydrolyzed into shorter oligopeptides by
endopeptidases and then further broken down into amino acids
by peptidase. Proteins or peptides may be hydrolyzed directly by
carboxypeptidases or aminopeptidases from the carboxyl or
amino ends, respectively, into shorter peptides or amino acids
which support protein synthesis in endosperm and embryo and so
proceeding of germination process.

• Lipids:
• The main storage oil in seeds is triacylglycerol stored in the oil
body (oleosomes), where it is hydrolyzed into fatty acids and
glycerol by lipases. The fatty acids are transported into
glyoxysomes to be degraded by β-oxidation, which is coupled with
the glyoxylate cycle to yield succinate. The synthesized succinate
is transported into the mitochondria to enter the TCA cycle to
form malate or oxaloacetate, which are transported into the
cytosol to produce sucrose

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• Phase III
• Protrusion of the radicle and continuous subsequent emergence of the
plumule occurs.

• In summary, water absorption by quiescent dry seeds activates many


kinds of hydrolysis and synthesis. The stored starch, proteins, oil, and
other complex nutrients in endosperm are hydrolyzed to sugars,
amino acids, and organic acids, which are absorbed by the scutellum
and transported to support the growth of the embryonic axis.

CHEESE PRODUCTION
• Cheese is produced from milk which cantains
carbohydrates (mainly lactose), proteins (mainly casein)
and fat.
• During cheese production, cheese making bacteria is
added to the milk to ferment it.
• The bacteria metabolise the lactose by breaking the it
into its constituent monomeric carbohydrates, glucose
and galactose by a enzyme β-galactosidase.
• In some cheese-making bacteria (Lactobacitlus
bulgaricus and most strains of Streptococcus
thermophiles), only the glucose is metabolised via
glycolysis to lactic acid while the galactose is exported
back into the milk, leading to the accumulation of
galactose. The galactose can react with protein in the
cheese during subsequent heating or storage leading to
the maillard reaction and non-enzymatic browning of
the cheese prepared using these bacteria.
• In some other cheese-making bacteria (Lactobacillus
plantarum, Lactobacillus belveticus can ferment both
the glucose and galactose in the milk and do not lead to
accumulation of galactose in the milk. Cheese prepared
using these bacteria hardly turn brown during storage.

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CARBOHYDRATE METABOLISM IN CHEESE-MAKING

Pathways for cheese bacteria that can


Pathways for cheese bacteria that can metabolise both
metabolise only glucose but not
glucose and galactose
galactose

METABOLIC GENERATION OF LACTATE IN CHEESE PREPARATION

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FURTHER READING
• Iskandar, C. F., Cailliez-Grimal, C., Borges, F., & Revol-Junelles, A. M.
(2019). Review of lactose and galactose metabolism in Lactic Acid
Bacteria dedicated to expert genomic annotation. Trends in food
science & technology, 88, 121-132.
• Ma, Z., Bykova, N. V., & Igamberdiev, A. U. (2017). Cell signaling
mechanisms and metabolic regulation of germination and dormancy
in barley seeds. The Crop Journal, 5(6), 459-477.

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