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HORT 604
APPLIED PHYSIOLOGY OF
HORTICULTURAL CROPS
Colegio de Postgraduados
Campus Montecillo
Summer 2007
Table of Contents
Topics Page
Growth Kinetics 38
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
root
stem
leaf
flower
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!
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.
STEM ANATOMY
Herbaceous Dicot or Gymnosperm - Primary Growth
STEM ANATOMY
Woody Dicot or Gymnosperm - Secondary Growth
STEM ANATOMY
Herbaceous Monocot - Primary Growth
ROOT ANATOMY
Herbaceous Dicot, Gymnosperm or Monocot - Primary Growth
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.
LEAF ANATOMY
Dicot
Monocot
(Similar to dicot, except no palisade, mesophyll is all spongy parenchyma)
LEAF ANATOMY
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
FLOWER STRUCTURE
FRUIT STRUCTURE
Example of a dry fruit Example of a fleshy fruit
SEED STRUCTURE
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.
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.
1) Auxin
2) Cytokinin
3) Gibberellic Acid
4) Ethylene
5) Abscisic Acid
6) Brassinosteroid
7) Jasmonic Acid
8) Salicylic Acid
9) Polyamines
AUXIN
SYNTHESIS
TRANSPORT
EFFECTS
4) Apical dominance - determined by correlative inhibition of apical bud, partly due to auxin
produced
5) Sprout Inhibitors – retard basal branching.
CYTOKININ
benzyladenine (BA)
zeatin root tips,
pyranylbenzyladenine
kinetin (not in plants) embryos
(PBA)
SYNTHESIS
adenine → zeatin
TRANSPORT
EFFECTS
shoot tips,
over 50 root tips,
(named by none embryos
consecutive
numbers)
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
4) Height control
• GA used to increase height
• growth retardants used to decrease height
ETHYLENE
SYNTHESIS
ETHYLENE INHIBITORS
TRANSPORT
EFFECTS
3) Respiration - increases
13) Leaf epinasty (curling and contortion or leaves) - causes in some plants
SYNTHESIS
EFFECTS
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.
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.
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.
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.
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.
Finger Printing
Gel Showing Banding from use of Different Restriction Enzymes
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.
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).
vivipary - germination of seeds inside the fruit while still attached to the parent plant.
1st Stage
2nd Stage
b) translocation to embryo
3rd Stage
SEED DORMANCY
1) Dry Seeds:
3) Embryo Rest:
4) Double Dormancy:
5) Chemical Inhibitors:
7) Light Requirement
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.
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.
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)
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
b) over any time LAR = leaf area2 - leaf area1 = LA2 -LA1
interval plant dry weight2 - plant dry weight1 W2 - W1
LAR is an indication of the efficiency of a given leaf area to produce a given plant size.
= 1 • ln W2 - ln W1
LA2 - LA1 t2 - t1
W2 - W1
NAR measures the accumulation of plant dry weight per unit leaf area per unit time.
It is a measure of efficiency of production.
CGR measures the efficiency of production of a total field of plants over a given soil area.
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.
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
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.
CARBON MOBILIZATION
Redistribution Between Sources and Sinks
(Fig. 10.19 from Taiz and Zeiger 1998, Fig. 3.61 from Larcher 1980)
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)
NUTRIENT MOBILITY
Immobile nutrients – deficiencies typically appear on newer growth and shoot tips first
MONOCARPIC SENESCENCE
Changing Sources and Sinks During Vegetative and Reproductive Growth
(Fig. 1 from Egli and Leggert 1973, Fig. 3 from Harper 1971)
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
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.).
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.
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.
1) Climacteric Fruit – a fruit where ethylene triggers an increase in respiration and the
ripening process..
3) Flower Senescence
4) Leaf Senescence
1) Ethylene
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
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..
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.
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.
mango (Fla.
14-21 little or no benefit >50 3.5 x
varieties)
12+ (slight
papaya (Solo) 12 28 2.3 x
benefit)
strawberry 7 7+ (off-flavor) 21 3x
14+ (slight
cucumber 14-Sep 49 3.5 x
benefit)
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)