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HORT 604
APPLIED PHYSIOLOGY OF
HORTICULTURAL CROPS

David Wm. Reed


Department of Horticultural Sciences
Texas A&M University

Colegio de Postgraduados
Campus Montecillo

Summer 2007

David Wm. Reed, Texas A&M University


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Table of Contents

Topics Page

Plant Anatomy and Morphology 2

Hormones and Elicitor Molelules 15

The Genetic Basis of Life 24

Genetically Modified Organisms (GMOs) or Transgenic Crops 30

Seed Germination, Dormancy and Priming 33

Growth Kinetics 38

Source Sink Relations 44

Senescence and Post Harvest Storage 50

David Wm. Reed, Texas A&M University


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Plant Anatomy and Morphology


A horticulturist who does not know the basic anatomy of plants is like is like a nurse that does
not know basic human anatomy. It could turn out to be down right uncomfortable where he/she
sticks that thermometer! So we are going to take a tour of plant structure. A working knowledge
of plant anatomy is absolutely essential in:
ƒ plant propagation: grafting, budding, division, cuttings, layering, tissue culture
ƒ pruning
ƒ making crosses in plant breeding
ƒ diagnosing plant disorders

Anatomy is very simply. Anatomists simply look at the outside and inside of plants and when
they see distinctive structures they give them a name. At the whole plant level, plants are
divided into four organs: The root, stem and leaf are vegetative organs, and the flower, and
resultant fruit, is a reproductive organ.
Plant Organs
ƒ root
ƒ stem
ƒ leaf
ƒ flower

Each organ is composed of three tissue systems:


Tissue Systems
ƒ dermal tissue system
ƒ vascular tissue system
ƒ ground or fundamental tissue system

Each tissue system is composed of distinctive tissues (epidermis, periderm, xylem, phloem,
cortex, pith and mesophyll), and tissues are in-turn composed of cells (parenchyma,
collenchyma, sclerenchyma, and specialized cells such as trichomes, vessels, companion cells,
laticifers, etc.).

Plants produce all these structures by growing from discrete clusters of dividing cells called
meristems. Herbaceous tissue is growth in length from: 1) apical meristems, which occur at
the end of every shoot and root, and 2) intercalary meristem at the base of grass leaves. Woody
tissue is due to growth in diameter from: 1) vascular cambium, which produce secondary
xylem (wood) and phloem, and 2) phellogen, which produces the periderm (bark).

Virtually all of the crops we grow in horticulture are monocots (linear leaves, ex. grasses, corn,
dracaena, and palm), dicots (broad-leaved plants, ex. oak, lettuce, apple) or gymnosperms
(leaves as needles and scales, ex. pine, juniper). The internal anatomy of monocots, dicots and
gymnosperms are sometimes similar and sometimes different. Different types of plants are not
like animals - all the tissues and organs are not always in the same location. Thus, one must
know the basic anatomical similarities and differences of each, or else you are not going to know
where to insert that thermometer - ouch!

David Wm. Reed, Texas A&M University


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ORGANS AND TISSUE SYSTEMS


Plants are composed of 3 vegetative organs and 1 reproductive organ.
Three tissue systems comprise each organ and are contiguous between each of the four organs.

David Wm. Reed, Texas A&M University


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HOW DO PLANTS GROW?

Meristems and Growth

Primary Growth - growth in length that gives rise to primary (herbaceous) tissues called the
primary plant body.

2 -Types
apical meristem or apex - the growing points located at the tips of stems and roots
intercalary meristem - the growth region at the base of grass leaves which causes
leaves to elongate.

Secondary Growth - growth in width or diameter which gives rise to secondary (woody
or corky) tissues called the secondary plant body.

lateral meristem - meristematic regions along the sides of stems and roots.

2 Types
vascular cambium or cambium - gives rise to secondary xylem (wood) on the
inside and phloem on the outside.

cork cambium or phellogen - gives rise to the periderm (bark).

David Wm. Reed, Texas A&M University


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STEM ANATOMY
Herbaceous Dicot or Gymnosperm - Primary Growth

(Fig. 16.1 from Esau 1960)

David Wm. Reed, Texas A&M University


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STEM ANATOMY
Woody Dicot or Gymnosperm - Secondary Growth

(Plate 28 from Esau 1965)

David Wm. Reed, Texas A&M University


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STEM ANATOMY
Herbaceous Monocot - Primary Growth

(Plate 58 from Esau 1965, Fig. 17.8 from Esau 1960)

David Wm. Reed, Texas A&M University


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ROOT ANATOMY
Herbaceous Dicot, Gymnosperm or Monocot - Primary Growth

(Plate 84 & 86 from Esau 1965)

David Wm. Reed, Texas A&M University


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ROOT ANATOMY
Woody Dicot or Gymnosperm - Secondary Growth

A woody dicot or gymnosperm root in secondary growth looks very similar to a stem in
secondary growth. The tissue is more porous and less dense, and the periderm is thinner. Rings
of xylem growth may not be as distinctive as occurs in stems. This is because roots of temperate
plants do not posses a distinctive “rest” or “physiological dormancy” period during the winter as
do buds and shoots. Root growth may occur whenever the soil moisture, fertility and
temperature are favorable.

(Fig.15.4 from Esau 1960)

David Wm. Reed, Texas A&M University


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LEAF ANATOMY

Dicot

(Plate 73 from Esau 1965)

Monocot
(Similar to dicot, except no palisade, mesophyll is all spongy parenchyma)

(Fig. 19.6 from Esau 1960)

David Wm. Reed, Texas A&M University


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LEAF ANATOMY

Gymnosperm

(Plate 78 from Esau 1965)

David Wm. Reed, Texas A&M University


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SUMMARY OF ANATOMY – VEGETATIVE STRUCTURES


MONOCOT DICOT GYMNOSPERM

PRIMARY
(herbaceous)
GROWTH

STEM

SECONDARY none
(woody)
GROWTH

PRIMARY
(herbaceous)
GROWTH
ROOT

SECONDARY
(woody) none
GROWTH

PRIMARY
(herbaceous)
GROWTH

LEAF

SECONDARY
(woody) none none none
GROWTH

David Wm. Reed, Texas A&M University


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FLOWER STRUCTURE

FRUIT STRUCTURE
Example of a dry fruit Example of a fleshy fruit

SEED STRUCTURE

David Wm. Reed, Texas A&M University


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Anatomical Structure and Function


"Structure and function" is a term used when the anatomy of a plant part explains how it
functions. Structure and function brings anatomy to the real world, and it is what makes anatomy
exciting. We are going to take a close look at one of the most important structure function
relationships in plants - translocation. The tissues responsible for long distance translocation in
plants are xylem and phloem.

Xylem is composed dead, hollow cells with perforated walls. The xylem cells are called vessel
elements or tracheids. . They are connected end to end and clustered side by side. They are like
a cluster of leaky pipes with holes on all sides. If you took sewer drain field pipe and connected
them end to end, and bundled many of them together side by side, you would have a perfect
model of xylem. Xylem only flows up. All xylem is dead and the water is "passively" pulled up
stems by transpiration of water from the leaves. It is like sucking water up a straw. In young
tissue, these bundles of xylem cells occur inside the vascular bundles, which are the stringy
tissue in herbaceous tissue (ex. veins in leaves). In woody plants, xylem is the wood. The
sapwood is functional because the hollow xylem cells are open and water easily flows up the
tubes. All the water flows up the sapwood. The heartwood is old clogged xylem, and does not
translocate water, and thus is not functional. The heartwood is clogged with resins and tannins
and this makes the heartwood both waterproof and prevents it from rotting.

Phloem is composed of specialized cells that remain alive and "actively" translocate solutes
(salts, sugars, metabolites, hormones, etc.) around plants. The phloem tissue is very
concentrated in sugars, amino acids, and many nutrients. It is the phloem that sucking insect,
such as aphids, puncture in order to feed on the sugar and nutrients... This is similar to a
mosquito piercing your veins and arteries as a food source.

Phloem flows both up and down and all around. It is commonly stated that phloem flows down,
but this is wrong. Phloem flows to where it is needed. Phloem flows from sources to sinks,
which will be discussed next.

David Wm. Reed, Texas A&M University


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HORMONES AND ELICITOR


MOLECULES
Hormone - an endogenous or naturally-occurring compound that is produced or
synthesized in one part of the plant and causes a change in physiology,
growth or development in another part of the plant; usually present in very
small quantities.

Elicitor Molecule - a compound which, when introduced in small concentrations to a living


cell system, initiates or improves the biosynthesis of specific compounds;
a compound with hormone-like activity.

Growth Substance - all naturally-occurring or synthetically produced compounds that affect the
physiology, growth and development of plants.

References
Moore, T.C. 1979. Biochemistry and Physiology of Plant Hormones. Springer-Verlag, NY.

Plant Hormones and Elicitor Molecules


Classically, plants have been known to contain five hormones, which are auxin, cytokinin,
gibberellic acid, ethylene and abscisic acid. Recently, other endogenous compounds have been
shown to elicit hormone-like reactions, which are brassinosteroids, jasmonic acid, salicylic acid
and polyamines. Some do not elevate these to the status of one of the five classical hormones, so
often they are called elicitor molecules.

1) Auxin

2) Cytokinin

3) Gibberellic Acid

4) Ethylene

5) Abscisic Acid

6) Brassinosteroid

7) Jasmonic Acid

8) Salicylic Acid

9) Polyamines

David Wm. Reed, Texas A&M University


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AUXIN

Naturally- Synthetic Structure Site of


Occurring Production
indolebutyric acid (IBA)
indoleacetic acid naphthaleneacetic acid shoot tips,
(IAA) (NAA) embryos
2,4-dichlorophenoxy-acetic
acid (2,4-D)

SYNTHESIS

tryptophan → indoleacetic acid

TRANSPORT

• 3:1 basipetal transport


• primarily in phloem parenchyma

EFFECTS

1) Cell elongation - causes acid induced cell wall growth

2) Cell division - stimulates

3) Tropism - response of plants to environmental or physical stimuli.


a) phototropism - response to light
b) geotropism - response to gravity
c) thigmotropism - response to touch

4) Apical dominance - determined by correlative inhibition of apical bud, partly due to auxin
produced
5) Sprout Inhibitors – retard basal branching.

6) Branch angle - causes wide branch angles

7) Fruit set - low concentrations stimulate

8) Fruit or flower thinning - high concentrations cause

9) Herbicides - 2,4-D at high concentrations

10) Adventitious root formation - a) stem and leaf cuttings


b) tissue culture

David Wm. Reed, Texas A&M University


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CYTOKININ

Naturally- Synthetic Structure Site of


Occurring Production

benzyladenine (BA)
zeatin root tips,
pyranylbenzyladenine
kinetin (not in plants) embryos
(PBA)

SYNTHESIS

adenine → zeatin

TRANSPORT

• xylem transported, found in root exudates


• primarily acropetal, but not necessarily polar

EFFECTS

1) Cell division - stimulates cell division; named after cytokinesis

2) Nutrient mobilization - nutrients transported towards high cytokinin concentration.

3) Apical dominance - high cytokinin/low auxin may overcome apical dominance

4) Chlorophyll breakdown - decreases chlorophyll breakdown

5) Leaf Aging or abscission - may delay

6) Seed germination - may overcome dormancy or stimulate germination

7) Adventitious shoot formation - a) leaf and root cuttings


b) tissue culture

8) Root growth - may be inhibitory to root growth

David Wm. Reed, Texas A&M University


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GIBBERELLIC ACID (GA)

Naturally- Synthetic Structure Site of


Occurring Production

shoot tips,
over 50 root tips,
(named by none embryos
consecutive
numbers)

SYNTHESIS (see next page)


mevalonate → farnesyl pyrophosphate →

→ geranylgeranyl pyrophosphate → copalyl pyrophosphate→ kaurene → GA

growth retardants - chemicals that block synthesis of GA; most block the ring closure steps
between geranylgeranyl pryophosphate → copalyl pyrophosphate →
kaurene.

TRANSPORT

• no polarity
• in phloem or xylem

EFFECTS

1) Protein synthesis - triggers de novo synthesis of some proteins, ex. α-amylase.

2) Cell elongation - primary stimulus for cell elongation

3) Rosette or dwarf plants - lack of endogenous GA often contributes to decreased height.

4) Height control
• GA used to increase height
• growth retardants used to decrease height

5) Flowering - may cause bolting in biennials

6) Fruit size - increases size of seedless grapes

7) Bud dormancy - may overcome and substitute for cold treatment

8) Seed germination - may increase or speed up

9) Sex expression - favors staminate flower formation on monoecious plants

David Wm. Reed, Texas A&M University


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Biosynthetic Pathway of Gibberellic Acid


(from Moore, 1979)

Mode of Action of Growth Retardants

• block ring closure between geranylgeranyl pyrophosphate and copalyl pyrophosphate


• block ring closure between copalyl pyrophosphate and kaurene

David Wm. Reed, Texas A&M University


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ETHYLENE

Naturally- Synthetic Structure Site of


Occurring Production

ethephon or ripening fruits,


ethylene
ethrel aging flowers,
(release ethylene germinating
inside plant) seeds,
wounded tissue

SYNTHESIS

methionine → s-adenosylmethionine → 1-aminocyclopropane-1-carboxylic acid → ethylene


(SAM) (ACC)

ETHYLENE INHIBITORS

ethylene inhibitors - chemicals that inhibit the synthesis or action of ethylene

Synthesis Inhibitors (block synthesis of SAM → ACC)


• AVG - aminoethoxyvinyl glycine
• MVG - methoxyvinyl glycine
• AOA - aminoacetic acid

Action Blockers (ethylene → block action)


• STS - silver thiosulfate
• CO2 - carbon dioxide
• Ni - nickel
• Co – cobalt
• MCP – 1-mehtylcyclopropane
o it is a gas that can saturate the receptor sites, and block action for several days
o EthylBloc – commercial compound

TRANSPORT

• diffusion as a gas throughout plant (in and out)

David Wm. Reed, Texas A&M University


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EFFECTS

1) Auxin transport - alters basipetal transport

2) Membrane permeability - increases

3) Respiration - increases

4) Cell elongation – decreases

5) Aerenchyma formation – induces aerenchyma formation under anaerobic or hypoxic


conditions (i.e. under low oxygen or flooded conditions)

6) Fruit ripening - stimulates in many fruits, ex. banana

7) Flowering - triggers flowering in some bromeliads, ex. pineapple

8) Flower fading - increases

9) Flower longevity - causes senescence (death) of cut flowers

10) Fruit color - decreases green, increases other colors

11) Seed germination - increases in some seeds

12) Leaf abscission (leaf drop) - causes in some plants

13) Leaf epinasty (curling and contortion or leaves) - causes in some plants

14) Sex expression - favors pistillate flower formation on monoecious plants

David Wm. Reed, Texas A&M University


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ABSCISIC ACID (ABA)

Naturally- Synthetic Structure Site of


Occurring Production

abscisic acid none


plastids,
especially
chloroplast

Historically also called:

abscisin - because early investigators found caused leaf abscission

dormin - because early investigators found caused dormancy

SYNTHESIS

mevalonate → farnesyl pyrophosphate → ABA

EFFECTS

1) Dormancy - causes bud or seed dormancy

2) Leaf abscission (leaf drop) - may cause in some plants

3) Stoma - causes stomata to close (a response to drought stress)

David Wm. Reed, Texas A&M University


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ELICITOR MOLECULES

Brassinosteroid
Effects:
• pollen tube growth
• stem elongation
• unrolling/bending grass leaves
• orientation of cellulose microfibrils
• enhanced ethylene production

Jasmonic Acid
Effects:
• defense mechanisms, promotes antifungal proteins
• growth inhibitor
• inhibit seed and pollen germination
• promotes curling of tendrils
• induces fruit ripening

Salicylic Acid
Effects:
• blocks ethylene synthesis
• induces flowering in some long day plants
• induces thermogenesis in voodoo lily
• defense mechanisms, promotes antifungal proteins

Polyamines
Effects:
• elicit cell division, tuber formation, root initiation, embryogenesis, flower development
and fruit ripening
• may not have a truly hormonal role; rather participate in key metabolic pathways
essential for cellular functioning.

David Wm. Reed, Texas A&M University


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THE GENETIC BASIS OF LIFE


(From http://generalhorticulture.tamu.edu)

Analogy

DEFINITIONS
DNA (deoxyribonucleic acid)- a double helix chain of sugar-phosphates (deoxyribo
sugar-phosphates) connected by nucleic acids (adenine, thymine, guanine, cytosine).
RNA (ribonucleic acid) - a single stranded chain of sugar-phosphates (ribo sugar-phosphates)
containing nucleic acids (adenine, uracil, guanine, cytosine).
nucleic Acids - organic acids that form the base pairs of DNA and single-bases of RNA.
Base Pairing of Nucleic Acids between the double strands of DNA
A- T (adenine-thymine)
G - C (guanine-cytosine)
Base Pairing of Nucleic Acids between DNA strands and RNA strands
A - U (adenine-uracil)
G - C (guanine-cytosine)
gene - a length of DNA that codes for the production of a protein or protein subunit.
- also codes for active RNAs (such as tRNA).
protein - a polymer or chain of amino acids.
enzyme - a protein that acts as a metabolic catalyst.

David Wm. Reed, Texas A&M University


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The Central Dogma of Molecular Biology


(From: http://www.accessexcellence.org)

Legend:
Transcription of DNA to RNA to protein: This dogma forms the backbone of molecular
biology and is represented by four major stages.
1. The DNA replicates its information in a process that involves many enzymes:
replication.
2. The DNA codes for the production of messenger RNA (mRNA) during transcription.
3. In eucaryotic cells, the mRNA is processed (essentially by splicing) and migrates from
the nucleus to the cytoplasm.
4. Messenger RNA carries coded information to ribosomes. The ribosomes "read" this
information and use it for protein synthesis. This process is called translation.
Proteins do not code for the production of protein, RNA or DNA.
They are involved in almost all biological activities, structural or enzymatic.

David Wm. Reed, Texas A&M University


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The Genetic Code


(From: http://www.accessexcellence.org)

DNA transfers information to mRNA in the form of a code defined by a sequence of


nucleotides bases. During protein synthesis, ribosomes move along the mRNA molecule
and "read" its sequence three nucleotides at a time (codon) from the 5' end to the 3' end.
Each amino acid is specified by the mRNA's codon, and then pairs with a sequence of three
complementary nucleotides carried by a particular tRNA (anticodon).
Since RNA is constructed from four types of nucleotides, there are 64 possible triplet
sequences or codons (4x4x4). Three of these possible codons specify the termination of the
polypeptide chain. They are called "stop codons". That leaves 61 codons to specify only 20
different amino acids. Therefore, most of the amino acids are represented by more than one
codon. The genetic code is said to be degenerate.

David Wm. Reed, Texas A&M University


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Restriction Enzymes Cut DNA at Specific Sequences


to Create “Sticky Ends”
(From: http://www.accessexcellence.org)

The EcoRI restriction enzyme--the first restriction enzyme isolated from E. Coli bacteria--is
able to recognize the base sequence 5' GAATTC 3'. Restriction enzymes cut each strand of
DNA between the G and the A in this sequence. This leaves "sticky ends" or single stranded
overhangs of DNA. Each single stranded overhang has the sequence 5" AATT 3'. These
overhanging ends will bond to a fragment of DNA which has the complementary sequence
of bases. See text of Background Paper for additional details.

David Wm. Reed, Texas A&M University


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Cloning DNA into a Plasmid to Produce Recombinant DNA


(From: http://www.accessexcellence.org)

Process by which a plasmid is used to import recombinant DNA into a host cell for cloning.
The plasmid carrying genes for antibiotic resistance, and a DNA strand, which contains the
gene of interest, are both cut with the same restriction endonuclease. They have
complementary "sticky ends." The opened plasmid and the freed gene are mixed with DNA
ligase, which reforms the two pieces as recombinant DNA. This produces recombinant
Deaths recombinant DNA stew transforms a bacterial culture, which is then exposed to
antibiotics. All the cells except those which have been encoded by the plasmid DNA
recombinant are killed, leaving a cell culture containing the desired recombinant DNA.
DNA cloning allows a copy of any specific part of a DNA (or RNA) sequence to be selected
among many others and produced in an unlimited amount. This technique is the first stage
of most of the genetic engineering experiments: production of DNA libraries, PCR, DNA
sequencing, et al.

David Wm. Reed, Texas A&M University


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Using restriction enzymes for Mapping or Finger Printing


When DNA from the same source is digested with a particular restriction enzyme it will always
give a set of the same sized fragments. For example if lambda bacteriophage DNA is cut with
EcoR1 we know that it will give six fragments of the sizes: 21.23, 7.42, 5.8, 5.65, 4.87, 3.53
kbp. This is because, mutations apart, the phage sequence will always be the same, and so EcoR1
cutting sites will always be present in the same places. The fragments can be separated and their
sizes determined by agarose gel electrophoresis.
We can use the positions of restriction enzyme sites as convenient markers along DNA
sequences. The map obtained can be used for DNA identification and to plan DNA
manipulations.

Finger Printing
Gel Showing Banding from use of Different Restriction Enzymes

David Wm. Reed, Texas A&M University


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Genetically Modified Organisms (GMO)


or Transgenic Crops
(From: http://www.colostate.edu/programs/lifesciences/TransgenicCrops/index.html)
Authors: Pat Byrne, Sarah Ward, Judy Harrington, Lacy Fuller (Web Master)

Crops and acreage

Transgenic crop production area by country (source: James, 2000b)


Area planted in 2000
Country Crops grown
(millions of acres)
soybean, corn, cotton,
USA 74.8
canola
Argentina 24.7 soybean, corn, cotton
Canada 7.4 soybean, corn, canola
China 1.2 cotton
South Africa 0.5 corn, cotton
Australia 0.4 cotton
Mexico minor cotton
Bulgaria minor corn
Romania minor soybean, potato
Spain minor corn
Germany minor corn
France minor corn
Uruguay minor soybean

David Wm. Reed, Texas A&M University


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Widely Used GMOs


Worldwide production area of transgenic crops – Traits
(source: Science 286:1663, 1999).
Area planted in 1999 (millions
Trait
of acres)
Herbicide tolerance 69.4
Bt insect resistance 22.0
Bt + herbicide tolerance 7.2
Virus resistance 0.3
Herbicide Tolerance Herbicide tolerant crops resolve many of those problems because they
include transgenes providing tolerance to the herbicides Roundup® (chemical name:
glyphosate) or Liberty® (glufosinate). These herbicides are broad-spectrum, meaning that they
kill nearly all kinds of plants except those that have the tolerance gene. Thus, a farmer can apply
a single herbicide to his fields of herbicide tolerant crops, and he can use Roundup and Liberty
effectively at most crop growth stages as needed.

Weed-
infested
soybean
plot (left)
and
Roundup
Ready®
soybeans
after
Roundup
treatment.
Source:
Monsanto

Bt Insect-Resistant Crops
"Bt" is short for Bacillus thuringiensis, a soil bacterium whose spores contain a crystalline (Cry)
protein. In the insect gut, the protein breaks down to release a toxin, known as a delta-endotoxin.
This toxin binds to and creates pores in the intestinal lining, resulting in ion imbalance, paralysis
of the digestive system, and after a few days, insect death.

European corn borer (left)


and cotton bollworm (right)
are two pests controlled by Bt
corn and cotton, respectively.
Source: USDA.

David Wm. Reed, Texas A&M University


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Bt insect-resistant crops currently on the market include


• Corn: primarily for control of European corn borer, but also corn earworm and
Southwestern corn borer. Cotton: for control of tobacco budworm and cotton bollworm
• Potato: for control of Colorado potato beetle. Bt potato has been discontinued as a
commercial product.

Papaya ringspot virus


Papaya is a tropical fruit rich in Vitamins A and C, but susceptible to a number of serious pests
and diseases. The transgenic variety UH Rainbow, resistant to the papaya ringspot virus, is
currently in production in Hawaii.

Papaya is an important source of


vitamins in tropical areas. Source:
USDA

Risks And Concerns


(http://www.colostate.edu/programs/lifesciences/TransgenicCrops/risks.html)
The introduction of transgenic crops and foods into the existing food production system has
generated a number of questions about possible negative consequences. People with concerns
about this technology have reacted in many ways, from participating in letter-writing campaigns
to demonstrating in the streets to vandalizing institutions where transgenic research is being
conducted. What are the main concerns? What scientific support is there for these concerns?
• damage to human health
allergenicity
horizontal transfer and antibiotic resistance
eating foreign DNA
cauliflower mosaic virus promoter
changed nutrient levels
• damage to the natural environment
Monarch butterfly
crop-to-weed gene flow
antibiotic resistance
leakage of GM proteins into soil
reductions in pesticide spraying: are they real?
• disruption of current practices of farming and food production in developed countries
crop-to-crop gene flow
• disruption of traditional practices and economies in less developed countries.

David Wm. Reed, Texas A&M University


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Seed
Germination, Dormancy and Priming
Terminology
pollination - deposition of pollen on the stigma of the pistil.

fertilization - the union of male and female gamete (nuclei, 1N) to produce zygote (2N).

double fertilization - in higher plants only (angiosperms)


- union of 1 1N male gamete with 1 1N female gamete (the egg) to
produce a 2N zygote; and union of 1 1N male gamete with 2 1N
polar nuclei to produce a 3N endosperm.

apomixis - development of an embryo without fertilization; hence, it is not true sexual


propagation even though it produces a seed.

parthenocarpy - development of fruit without seeds.

vivipary - germination of seeds inside the fruit while still attached to the parent plant.

Seed Dormancy Terminology


Primary Dormancy
Old Term New Term Definition
Quiescence Ecodormancy Dormancy imposed by an external unfavorable
environmental factor or external structure.

Example: too dry, external hard seed coat


Correlative Paradormancy Dormancy imposed by physiological factor external to
Inhibition the embryo.

Example: inhibitors in testa or pericarp


Rest Endodormancy Dormancy imposed by a physiological factor internal to
or the embryo.
Physiological
Dormancy Example: embryo rest
Secondary Dormancy
Photodormancy Dormancy due to lack of light (red) in light requiring
seeds.

David Wm. Reed, Texas A&M University


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STAGES OF SEED GERMINATION

1st Stage

a) imbibition - initial absorption of water to hydrate seed

b) activation of metabolism - increased respiration and protein synthesis

2nd Stage

a) digestion of stored food - for example, starch to sugars in cotyledon or endosperm

b) translocation to embryo

3rd Stage

a) cell division and continued growth and development of seedling

David Wm. Reed, Texas A&M University


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SEED DORMANCY

CAUSED BY TYPE DORMANCY HOW OVERCOME?

1) Dry Seeds:

dehydration of seed quiescence sow in moist environment

2) Seed Coat Dormancy or Hardseededness:

hard seed coat impermeable quiescence scarification - physical or


to water and gases chemical abrasion of seed coat.

3) Embryo Rest:

low growth promoters rest stratification - cold (35-40 oF


and/or high growth (or physiological moist storage for 4-12 weeks.
inhibitors in embryo dormancy)

4) Double Dormancy:

hard seed coat plus quiescence scarification then stratification


embryo rest and rest (or
physiological dormancy)

5) Chemical Inhibitors:

inhibitors in pericarp (fruit correlative I) remove fleshy pericarp (fruit


wall) or testa (seed coat) inhibition wall) or testa (seed coat).
2) leach in running water if
pericarp or testa is dry.
6) Immature Embryo:

underdeveloped or developmental I) after ripening - store for 4-6


rudimentary embryo dormancy weeks under ambient
conditions
2) warm stratification - warm
moist storage.
3) embryo culture

7) Light Requirement

phytochrome in Pr form secondary I) expose to any white light


dormancy 2) expose to red light
3) sow shallow or on surface

David Wm. Reed, Texas A&M University


36

Seed Treatments to Enhance Germination

Seed Priming
Seed priming is a seed treatment that imbibition and activation of the initial metabolic events
associated with seed germination, but prevents radicle emergence and growth. Obviously, seeds
are tolerant of desiccation, and even though during seed priming imbibition allows water uptake,
the tolerance to desiccation is not lost. Thus, the seed can be dried again and stored. If the seeds
are primed too long, desiccation tolerance will be lost, and the seeds may loose viability upon re-
drying. The secret to successful seed priming is to stop the priming treatment at just the right
time to allow re-drying.

The advantage of primed seed is that when the primed seeds are planted their germination is
faster and more uniform.

David Wm. Reed, Texas A&M University


37

Types of seed priming


1) Osmopriming (osmoconditioning):
This is the most common technique used. The seeds are soaked in an osmotic solution to
allow imbibition and metabolic activation, but the osmotic conditions do not allow expansion
and growth of cells. Osmotica used are: mannitol, polyethyleneglycol (PEG) or salts such
as KCl.

2) Hydropriming:
Imbibition is obtained by partially hydrated the seeds using a limited amount of water by
exposing them to a limited amount of water, using very humid air or exposing them for a
short time in warm water.

3) Matrix priming:
A solid, insoluble matrix is used to obtain a water solution with low water potential. The
matrix potential keeps the water potential low. Vermiculite, diatomaceous earth or cross-
linked highly water-absorbent polymers are used.

Hormone Treatments to Enhance Germination


Seeds of some species are very difficult and slow to germinate due to primary and secondary
dormancy, the need for after ripening periods, immature embryos, etc. Many of the seeds
respond to hormones to increase the speed of germination, uniformity of germination and/or
percent germination. For example, the seeds of many tropical foliage plants are difficult to
geminate, but respond to hormonal treatments. Hormones can also be added to seed priming
treatments.

1) cytokinin – 100 to 200 mg/liter for a 12-24 hour soak.

2) gibberellic acid – 200 to 1,000 mg/liter fro 12-24 hour soak

David Wm. Reed, Texas A&M University


38

GROWTH KINETICS
Growth - an irreversible increase in size, mass or number.
Many growth phenomena in nature exhibit a logarithmic or exponential increase. The size, mass
or number increases by a constant, similar to simple compound interest. The principal (current
size, mass or number) times the interest rate (growth rate) yields the interest (growth increase for
that day). The interest is added to the principal, to yield a new principal. The new principal
times the interest rate yields and even higher interest for the next day, which again is added back
to the principal. So growth occurs at a compounded rate (logarithmic or exponential growth).
Absolute Growth Rate (AGR)
If you plot growth (size, mass or number) versus time, a constantly increasing growth curve is
obtained. If you calculate the slope between any two times, you get the absolute growth rate,
which is the change in actual growth over time. You get a different slope, hence different AGR
for each pair of times chosen to calculate the slope. (Fig. 2.23A, Wareing and Philips 1981)

Relative Growth Rate (RGR)


If you plot the logarithm of growth (size, mass or number) versus time, a linear line is obtained.
If you calculate the slope of the line, you get the relative growth rate, which is the change in
relative growth over time. Since the line is linear, you get the same RGR, regardless of which
time interval chosen to calculate the slope. (Fig. 2.23A, Wareing and Philips 1981).

David Wm. Reed, Texas A&M University


39

GROWTH KINETICS- con't

Sigmoidal Growth Curve


Exponential growth can never be sustained indefinitely. Eventually, substrates are depleted, the
population exceeds the area available, tissues or individuals begin to die, etc., which decreases
the growth rate. Growth may still increase, but at a reduced rate (ex. if crowding causes
shading), it may reach a steady state (everything is in equilibrium, for example in a population),
or growth may begin to decrease (ex. due to death or senescence of individuals or plant parts). If
you plot long term growth versus time you get the classical sigmoidal growth curve. If you plot
the logarithm of the sigmoidal growth curve, you get a linear line during the exponential phase,
after which the curve decreases over time. (Fig. 2.24, Wareing and Philips 1981)

Changes In Growth Rates Over Time


If you calculate the absolute growth rate (AGR) over increments of time, then plot AGR versus
the time interval, you get a bell-shaped curve, i.e. the AGR changes constantly with time. If you
calculate the relative growth rate (RGR) over increments of time, then plot RGR versus the time
interval, you get a straight-line region during the logarithmic phase followed by a decreasing
RGR. The RGR is constant during the logarithmic phase. (Fig. 2.27, Wareing and Philips 1981).

David Wm. Reed, Texas A&M University


40

MATHEMATICAL MODELS OF GROWTH

Linear Model – Used During Logarithmic or Exponential Phase

ln n = ln no + (slope) (time)
where n = number, size (height, leaf area), or mass (dry weight, fresh weight) at any time > 0.
no = number, size (height, leaf area,), or mass (dry weight, fresh weight) at time = 0.
slope = rate of growth

Or more commonly expressed as the slope equation


y = a + bx

y = intercept + (slope) (x)

Absolute Growth Rate (AGR)


AGR = dn
dt

= n2 - n 1 yields average slope over that time interval


t2 - t 1

Relative Growth Rate (RGR)


RGR = dn • 1
dt n

= ln n2 - ln n1 yields constant slope during logarithmic phase


t2 - t 1

David Wm. Reed, Texas A&M University


41

QUANTITATIVE MEASUREMENTS OF GROWTH

Leaf Area Ratio (LAR)

a) over life of crop LAR = final leaf area = LA


final plant dry weight W

b) over any time LAR = leaf area2 - leaf area1 = LA2 -LA1
interval plant dry weight2 - plant dry weight1 W2 - W1

; units = cm2 g-1 or cm2/g

LAR is an indication of the efficiency of a given leaf area to produce a given plant size.

Net Assimilation Rate (NAR)

NAR = RGR = 1 • RGR


LAR LAR

= 1 • ln W2 - ln W1
LA2 - LA1 t2 - t1
W2 - W1

= W2 - W1 • ln W2 - ln W1 ; units = g cm-2 day-1 or g/cm2/day


LA2 - LA1 t2 - t1

NAR measures the accumulation of plant dry weight per unit leaf area per unit time.
It is a measure of efficiency of production.

Leaf Area Index (LAI)

LAI = leaf area = LA ; units = cm2leaf cm-2soil or cm2leaf/cm2soil


soil area A

Measures the fraction of crop cover.


LAI is near 0 at planting, and is usually 2-3 at full canopy coverage

Crop Growth Rate (CGR)

CGR = NAR • LAI ; units = g cm-2soil day-1 or g/cm2soil/day

CGR measures the efficiency of production of a total field of plants over a given soil area.

David Wm. Reed, Texas A&M University


42

APPLICATION OF QUANTITATIVE MEASUREMENTS OF GROWTH

Efficiency of Different Species of Plants


The following table gives the net assimilation rates (NAR) of various species. The higher the
NAR the more efficient the species, which usually translates into higher growth rates. (from
Table 3.10, Larcher 1980)

Net Assimilation Rate


(mg dry matter per dm2 leaf area per day)
Average Over During Main
Plant Type Growing Season Growth Phase
C4 Grasses >200 400-800
Herbaceous C3 Plants
Grasses 50-150 70-200
Dicots 50-100 100-600
Woody Dicots
Topical and Sub-Tropical 10-20 30-50
Deciduous Temperate Trees 10-15 30-100
Conifers 3-10 10-50
Ericaceous Shrubs 5-10 15
CAM Plants 2-4 10

Efficiency of Sun versus Shade Plants


The following table gives the net assimilation rates (NAR), leaf area ratio (LAR), and relative
growth rate (RGR) of shade versus sun plants at both high and low light intensities. (from Table
3.1, Leopold and Kreidmann1975).

Note: At low light intensities, the sun plant has 6-fold decrease in NAR and tries to compensate
by increasing its LAR (i.e. produces about 2-fold more and/or larger leaves), but the RGR still
decreases dramatically. At low light intensities, NAR of the shade plant only decreases 3-fold,
and increases its LAR 2.4 fold, both of which help maintain a higher RGR; in other words the
shade plants have adapted themselves to the lower light intensity.

NAR LAR RGR


2
% Daylight mg/cm / wk % cm2/g g/g wk %
Sun Plant - Sunflower
100% 8.0 100 82 0.66 100
24% 2.9 36 140 0.42 64
12% 1.3 17 170 0.23 35
Shade Plant - Impatiens
100% 6.1 100 132 0.80 100
24% 3.3 54 239 0.78 98
12% 2.0 33 315 0.63 79

David Wm. Reed, Texas A&M University


43

APPLICATION OF QUANTITATIVE MEASUREMENTS OF GROWTH - con't

Effect of Leaf Area Index (LAI) on


Net Assimilation Rate (NAR) and Crop Growth Rate (CGR)
Note that as the LAI increases (due to greater canopy coverage of soil), the NAR (productivity of
each plant) decreases (probably due to increased plant-plant shading), but the CGR (productivity
of the entire crop over a given area of soil) increases. Thus, the best LAI is somewhere around 4.
(from Fig. 3.64, Larcher 1980)

Use of Quantitative Growth Measurements to Explain


Other Growth Phenomena
Increasing ambient carbon dioxide increases photosynthesis, which in turn increases growth. In
tomato and bean, increasing carbon dioxide increases both total plant growth, as measured by
increased RGR, and the efficiency of growth, as measured by increased NAR. This increased
growth efficiency allows the plant to have a smaller shoot system (decreased LAR), which is the
source, while still enhancing the size of the root system (see increased root/shoot ratio), which is
a sink (from Table 3-2, Leopold and Kriedemann 1975).

Tomato Bean
300 ppm 1,000 ppm 300 ppm 1,000 ppm
CO2 CO2 CO2 CO2
RGR (mg g-1 d-1) 222 254 122 172
NAR (mg dm-2 d-1) 71 89 46 80
LAR (dm2 g-1) 3.0 2.8 3.2 2.7
root/shoot ratio 0.19 0.21 0.18 0.25

David Wm. Reed, Texas A&M University


44

Source Sink Relations


PHLOEM AND XYLEM TRANSLOCATION
(Figure 3.9 and Table 3.8 from Marshner 1986, Summary from Bidwell 1974)

Fig 3 9 Long-distance transport in xylem


(X) and phloem (P) in a stem with a
connected leaf, and xylem-to-phloem
transfer mediated by a transfer cell (T).

Table 3.8. Solutes in the Phloem and Xylem Exudates of tobacco.


T Phloem exudate Xylem
(stem incision) exudate
Concentrati
on ratio
a pH 7-8-8-0 (tracheal) phloem/
Dry matter (/i / l)*170-196 C H 5 6
M-1-25 C9 l
155-163C
C
Sucrose 155-168 ND —
Reducing sugars Absent NA —
Amino compounds 10,808-0 283-0 38-2
Nitrate ND NA —
Ammonium 45-3 9-7 4-7
Potassium 3,673-0 204-3 18-0
Phosphorus 434-6 68-1 6-4
Chloride 486-4 63-8 7-6
Sulfur 138-9 43-3 3-2
Calcium 83-3 189-2 0-44
Magnesium 104-3 33-8 3-1
Sodium 116-3 46-2 2-5
Iron 9-4 0-60 15-7
Zinc 15-9 1-47 10-8
Manganese 0-87 0-23 3-8
4
Copper 1-20 0-11 10-9
ND, Not detectable; NA, data not available., 'Milligrams per milliliter.

SUMMARY. The general conclusions about the pathways and tissues of translocation:
1. Salts and inorganic substances move upward in the xylem.
2. Salts and inorganic substances move downward in the phloem.
3. Organic substances move up and down in the phloem.
4. Organic nitrogen may move up in the xylem (trees) or phloem (herbaceous plants).
5. Organic compounds like sugar may be present in the xylem sap in large concentrations
during the spring when sap rises in trees before the leaves emerge.
6. Lateral translocation of solutes from one tissue to another occurs, presumably by
normal mechanisms of transfer (osmosis, active transport, and so on).
7. Exceptions to these generalizations are known to occur.

David Wm. Reed, Texas A&M University


45

CARBON MOBILIZATION
Redistribution Between Sources and Sinks
(Fig. 10.19 from Taiz and Zeiger 1998, Fig. 3.61 from Larcher 1980)

Figure 10.19 Autoradiographs of a leaf of summer squash (Cucurbita pepo), showing


the transition of the leaf from sink to source status. In each case, the leaf imported 14C
from the source leaf on the plant for 2 hours. Label is visible as black accumulations.
(A) The entire leaf is a sink, importing sugar from the source leaf. (B-D) The base is
still a sink. As the tip of the leaf loses the ability to unload and stops importing sugar,
as shown by the loss of black accumulations in B through D, it gains the ability to load
and to export sugar. (From Turgeon and Webb 1973, courtesy of R. Turgeon.)

Fig. 3.61. Variations in starch deposition by trees throughout the year. Maximal accumulation of
starch is indicated by black, large amounts by cross-hatching, and small amounts by stippling; in
the parts left white, starch is present in traces or not at all. Fagus sylvatica (Central Europe): 1, just
before leaf emergence in the spring; 2, during leaf unfolding; 3, midsummer; 4, just before abscission
in the autumn; 5, conversion of starch to soluble carbohydrates at low temperatures during winter.
After Fischer (1891), Gaumann (1935), and K. Kober (unpubl.). Abies veitchii (Japan): 1, during
growth of new shoots in spring; 2, late summer; 3, during winter frost. After Kimura (1969). Olea
europaea (Northern Italy): 1, during shooting and flowering in spring; 2, during a dry period in
midsummer; 3, in winter after the end of the rainy season. After Thomaser (1975). For the storage
dynamics of Atlantic dwarf shrubs see Stewart and Bannister (1973), and Grace and Woolhouse
(1973); of chaparral species, Mooney and Hays (1973); of mountain plants, Larcher (1977) and
Zachhuber and Larcher (1978)

David Wm. Reed, Texas A&M University


46

NUTRIENT MOBILITY

Redistribution Between Sources and Sinks


(Fig. 13-12 from Bidwell 1974, Table 3.9 from Marschner 1986)

Figure 13-12 (opposite). A sequence of six autoradiograms showing the fate of an


aliquot of 35S absorbed as 35S04 during a 1-hr absorption period. The plants, after the
hour in the nutrient solution containing the tracer, were removed to a normal
(nonradioactive) solution where they remained for the following periods: A, 0 hr;
B, 6 hr; C, 1 2 hr; D, 24 hr; E, 48 hr; and F, 96 hr. Most of the 35S, which moved
directly into the mature leaves, was withdrawn within 1 2-24 hr. It moved predomi-
nantly into younger leaves near the stem apex, where it remained. [From 0. Biddulph:
Plant Physio/. 33:295 (1958). Used with permission. Photograph courtesy Dr.
Biddulph.]

Table 3.9. Mobility of Mineral Elements in Phloem


Mobile Intermediate Immobile
Potassium Iron Manganese Lithium
Rubidium Zinc Copper Calcium
Sodium Molybdenum Strontium
Magnesium Barium
Phosphorus Boron
Sulfur
Chlorine

From Bukovac and Wittwer (1957).

David Wm. Reed, Texas A&M University


47

DIAGNOSING NUTRIENT DEFICIENCIES


Based on Nutrient Mobility
(from Vetanovetz 1996)

Mobile Nutrients – deficiencies typically appear on older growth first.

Immobile nutrients – deficiencies typically appear on newer growth and shoot tips first

David Wm. Reed, Texas A&M University


48

MONOCARPIC SENESCENCE
Changing Sources and Sinks During Vegetative and Reproductive Growth
(Fig. 1 from Egli and Leggert 1973, Fig. 3 from Harper 1971)

Fig. 1. Dry matter accumulation patterns for Kent and D66-


5566, 1971.

Fig. 3. Seasonal uptake and accumulation of N, P, K, Ca, and


Mg by soybeans at weekly intervals1 from field hydroponic
gravel culture systems.

David Wm. Reed, Texas A&M University


49

EPISODIC GROWTH OF TEMPERATE WOODY PLANTS


Cycling Between Shoot and Root Growth and Implications on Fertilizer Timing
(Fig. 2 on growth from Mertens and Wright 1978, Fig. 2 on uptake from Hershey and Paul 1983,
Table 1 from Gilliam and Wright 1978)

Fig. 2. Root and shoot growth rates of Fig. 2. Uptake rates for K+ and Mg 2+ for a
'Helleri' holly grown at 150 ppm N applied as single plant of Euonymus japonica (plant 5).
20N-8.7P-16.5K soluble fertilizer. Bars indicate periods of shoot elongation

Table 1. Effect of the time and no. of weekly fertilizer


applications during 1st growth flush on tissue N accumulation
and subsequent shoot and root dry wt of 'Helleri' holly.
Week No. %N Shoot dry Root dry
Fertilizer Appl wt (g) wt (g)
Applied
1 1 1.88 5.1 2.4
2 1 1.99 5.3 2.4
3 1 2.01 4.9 2.9
4 1 2.27 6.2 2.4
5 1 2.04 5.2 2.2
1-2 2 2.10 5.5 2.3
2-3 2 2.23 5.9 2.4
3-4 2 2.26 6.9 2.4
4-5 2 2.45 6.1 2.0
1-2-3 3 2.13 6.1 1.9
2-3-4 3 2.38 7.1 1.9
3-4-5 3 2.58 6.7 2.0
1-2-3-4 4 2.69 6-5 1.8
2-3-4-5 4 2.55 6.5 1.6
1-2-3-4-5 5 2.59 7.0 1.7

David Wm. Reed, Texas A&M University


50

Senescence and Post-Harvest Storage


MONOCARPIC SENESCENCE
Monocarpic senescence literally means “flower once then die”. During the reproductive phase,
the “sink” demand of the developing flowers, fruit then seed can drain the vegetative “sources”
to the point that senescence occurs.

Fig. 1. Dry matter accumulation patterns for Kent and D66-


5566, 1971. (from Egli and Leggert 1973)

Fig. 3. Seasonal uptake and accumulation of N, P, K, Ca, and


Mg by soybeans at weekly intervals1 from field hydroponic
gravel culture systems.(from Harper 1971)

David Wm. Reed, Texas A&M University


51

RESPIRATION AND SENESCENCE

All living organisms must conduct respiration in every living cell and at all times. Sometimes
respiration is very fast, for example if the organ if actively growing, and sometimes it barely
perceptible, for example if the organ is dormant. Respiration breaks down glucose and uses the
energy that was in the carbon-carbon bond to make metabolic energy (mainly a compound called
adenosine triphosphate or ATP). Carbon dioxide is given off as a by-product. If there is no
oxygen around, then only partial respiration occurs in the form of anaerobic fermentation. This
produces ethanol as a by-product and is the basis of wine making and all fermentation (yogurt,
cheese, etc.).

One process involving respiration that is particularly important to horticulturist is ripening of


fruit. In climacteric fruit, the respiration rises very rapidly during ripening, then decreases as
the fruit senesces. If you prevent or decrease the rise of respiration, then you can prolong post-
harvest storage life. Ethylene is what causes the increase in respiration, so decreasing ethylene
is also a strategy used to increase post-harvest storage life.

What are other ways to decrease respiration and prolong the storage life of fruit and vegetable
produce or cut flowers? Look at the equation for respiration. We can make the reaction go
slower by either decreasing things on the left side of the arrow or increasing things on the right
side of the arrow. Practically, we can decrease respiration by either increasing carbon dioxide
or decreasing oxygen. You want to increase carbon dioxide to about 2-5% (up from about 350
ppm in the ambient air) and/or decrease oxygen to about 3% (down from 21% in the ambient
air). You never want to decrease oxygen to near zero, because anaerobic fermentation would
occur and anaerobic bacteria might start growing.

Of course the easiest way to decrease respiration is to decrease temperature. You may not
have thought about it, but the refrigerator in your house is nothing more than a respiration
inhibitor chamber.

All of the above is the basis of controlled-atmosphere storage.

If in addition to the above, if you store produce or flowers under a light vacuum, you will pull
the ethylene out of the inside of the plant and the atmosphere around the plant. This will
dramatically decrease respiration. This is called hypobaric storage.

Summary, we can decrease respiration by doing the following:


ƒ decrease temperature
ƒ decrease oxygen
ƒ decrease pressure, e.g. light vacuum
ƒ decrease ethylene
ƒ increase carbon dioxide

David Wm. Reed, Texas A&M University


52

NET CHEMICAL EQUATION FOR RESPIRATION

BIOCHEMICAL REACTIONS OF RESPIRATION

David Wm. Reed, Texas A&M University


53

Ethylene, Respiration and Senescence Relations

1) Climacteric Fruit – a fruit where ethylene triggers an increase in respiration and the
ripening process..

Climacteric Fruit Ripening and Climacteric Rise

Climacteric Fruit Non-Climacteric Fruit


apple olive bell pepper
apricot pawpaw blueberry
avocado peach cherry
banana pear grape
cantaloupe plum pineapple
fig persimmon strawberry
guava ? tomato citrus
mango watermelon

2) Non-Climacteric Fruit color – used to cause degreening of citrus

3) Flower Senescence

a) Flower fading – flower fade after pollination

b) Flower longevity - causes senescence (death) of cut flowers

4) Leaf Senescence

a) Leaf epinasty (curling and contortion or leaves) - causes in some plants

b) Leaf abscission (leaf drop) - causes in some plants

David Wm. Reed, Texas A&M University


54

Manipulating Ethylene, Respiration and Senescence

1) Ethylene

Biosynthetic Pathway of Ethylene Synthesis


methionine → s-adenosylmethionine → 1-aminocyclopropane-1-carboxylic acid → ethylene
(SAM) (ACC)

Ethylene inhibitors - chemicals that inhibit the synthesis or action of ethylene

Ethylene Synthesis Inhibitors (block synthesis of SAM → ACC)


• AVG - aminoethoxyvinyl glycine
• MVG - methoxyvinyl glycine
• AOA - aminoacetic acid

Ethylene Action Blockers (ethylene → block action)


• STS - silver thiosulfate
• CO2 - carbon dioxide
• Ni - nickel
• Co – cobalt
• MCP – 1-mehtylcyclopropane
o it is a gas that can saturate the receptor sites, and block action for several days
o EthylBloc – commercial compound

2) Temperature
• respiration decreases when temperature decreases.
o
• respiration ceases at about freezing temperatures (32 F)
• increasing temperature increases respiration, until temperature gets too high, then
respiration decreases when tissue deteriorates

3) Oxygen
• respiration decreases when oxygen decreases
• under very low to no oxygen, anaerobic respiration occurs.

4) Carbon Dioxide
• respiration decreases when carbon dioxide increases

David Wm. Reed, Texas A&M University


55

Post-Harvest Storage to Extend Shelf Life


1) Refrigeration

• decrease temperature – mid-30’s oF

2) Controlled Atmosphere Storage – CA Storage

How Apples are Packaged and Stored


Apples are stored in cold storage warehouses. Inside a regular
warehouse, apples can be stored for about 5 months because it
is cooled to 30-32 degrees Fahrenheit.
Inside a special controlled atmosphere warehouse, apples can
be stored for almost 12 months because the temperature,
humidity, oxygen and carbon dioxide are constantly monitored
and controlled to prevent the fruit from ripening too quickly.
(from Dole: www.dole5aday.com/ReferenceCenter/
Encyclopedia/Apples/apple_transported2.jsp

• low temperature - mid-30’s oF


• high CO2 - approx. 2-5%)
• low 02 - approx. 3%)
• high humidity -approx. 90%)
• ethylene removed -scrubbed

3) Hypobaric Storage low pressure storage, i.e. a light vacuum.

Hypobaric Storage Shipping Container


(from http://www.refrigeratedvehicles.com/)

• same as above, plus


• low pressure
o decreases 02
o decreases ethylene

David Wm. Reed, Texas A&M University


56

4) Modified Atmosphere Packaging – MAP

MAP broccoli
(from http://www.packagingdigest.com/articles/200203/32.php)

MAP uses selectively permeable bags and the fruit or vegetable’s own respiration to
maintain an increased level of carbon dioxide and decreased level of oxygen, but avoiding
low enough oxygen to avoid anaerobic respiration.

Fruits and vegetables continue to respire after harvest. If you seal them in a plastic bag, the
produce will deplete the atmosphere in the bag of oxygen and will cause the produce to
undergo anaerobic respiration. This will causes ethanol and off-flavors to form and may
allow anaerobic bacteria to grow and cause spoilage.

In MAP, the produce is place in a selectively permeable bag that allows oxygen, carbon
dioxide and ethylene to diffuse in and out so equilibrium is set-up between the inside of the
bag to the outside of the bag. The goal is to use a bag that allows some oxygen to diffuse in
to avoid anaerobic fermentation, but allow excessive carbon dioxide and ethylene to escape..

Permeability of Various Films


Thickness Permeability (l/m2/d/atm)
Film
(micron) O2 CO2
polyvenylchloride 14-18 20 120
ethylenevenlyacetate 10-25 32 134
low density polyethylene 25-50 6 20
p0lystyrene 50 4 13

The bag must be designed for each fruit and vegetable. Produce with very high rates of
respiration require a bag that allows more oxygen to diffuse in to avoid anaerobic respiration.

David Wm. Reed, Texas A&M University


57

Respiration Rates of Vegetables


Class Respiration Commodities
Rate (mg / kg / hr)
Very Low Below 10 Onion
Low 10 - 20 Cabbage, tomato
Moderate 20 - 40 Carrot, celery
High 40 - 70 Lettuce, radish
Very High 70 - 100 Spinach, bean
Extremely High Above 100 Broccoli, pea

Two Types of MAP

1) Passive MAP

The produce is put in a bag. If the permeability of the bag is properly matched with the
respiration of the produce, the ideal atmosphere will evolve inside the sealed bag.
Absorbers may be added to scavenge ethylene.

2) Active MAP

The produce is put in a bag, and the air in the bag is replaced with air that has the proper
mixture of oxygen and carbon dioxide. Absorbers may be added to scavenge ethylene.

David Wm. Reed, Texas A&M University


58

Maximum Storage Time with Various Storage Methods


(from http://atlasuhv.com/products/hypobaric_storage/hypobaric_storage.php

Maximum Storage Time (days)


Hypobaric
Commodity Hypobaric Benefit
Standard Control
Advanced Factor
Refrigeration Atmosphere
Atmosphere
spinach 14-Oct slight benefit 50 5x

avocado (Lula) 30 42-60 >102 3.5 x

banana 14-21 42-56 150 11 x

cherry (sweet) 14-21 28-35 56-70 4x

juice loss, peel


lime (Persian) 14-28 90+ 6.5 x
thickens

mango (Fla.
14-21 little or no benefit >50 3.5 x
varieties)

12+ (slight
papaya (Solo) 12 28 2.3 x
benefit)

pear (Bartlett) 60 100 200 3.3 x

strawberry 7 7+ (off-flavor) 21 3x

slight benefit - off


asparagus 14-21 28-42 2x
odors

14+ (slight
cucumber 14-Sep 49 3.5 x
benefit)

green pepper 14-21 no benefit 50 3.5 x

mushroom 5 6 21 4.2 x

apples
200 300 300+ 1.5 x
(various)

carnation
21-42 no benefit 140 6.6 x
(flower)

protea (flower) <7 no benefit 30+ 4.2 x

rose (flower) 14-Jul no benefit 42 6x


The above data from S.P. Burg in Postharvest Physiology and Hypobaric Storage of
Fresh Produce, CABI Publishing, 2004, ISBN 0 85199 801 1

David Wm. Reed, Texas A&M University