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Weed biology

Gyombiolgiai s gyomszablyozsi
ismeretek modul
Main topics

The aim of the subject to train modern weed control


knowledge based on biological background.
Students must know:
Biological qualities of important weeds,
Ecological claims of important weeds
We consider stressed educational task forming of
integrated approach in weed control.

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Main topics

Definition, classification and adaptation of weeds


Plant life forms
Weed survey methods
Reproduction biology of annual weeds
Reproduction biology of perennial weeds
Competition
Allelopathy
Herbicide resistance of weeds

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Definition of weed

Weeds are the greatest biological challenge to


agricultural production, whether conventional or
organic!
What makes a plant a weed?
Cost of weeds
Why do weeds always win? Biology!
Get to know the enemy
Anatomy of a weed
Strategies for weed management

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Definition of weed

Blatchley 1912: A plant out of the place or growing


where it is not wanted.
Robbins et al. 1942: These obnoxious plants are
known as weeds.
The European Weed Research Society 1986: Any
plant or vegetation, excluding fungi, interfering with
the objectives or requirements of people.

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Definition of weed

Weed is a plant or plant parts (stolon, rhizome,


tuber, bulb etc.) in any growing stage, which is
growing where it is not desired.
Hunyadi (1974)

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Importance of weeds

There are about 200.200 plant species in the World, and


approximately 6700 species of weeds.
76 species belong to most dangerous weed species of the
World, but only 18 have prominent importance.

The most important families in


the world :

Graminaceae Cruciferae
Compositae} ~ 40% Fabaceae
Cyperaceae Convolvulaceae
Polygonaceae Euphorbiacea
Amarantaceae Chenopodiacea

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Importance of weeds

The most important weed familyes of the World

Family Number of species


1. Gramineae 44
38%
2. Compositae 32
3. Cyperaceae 12

4. Polygonaceae 8 65%
5. Amaranthaceae 7
6. Cruciferae 7

7. Fabaceae 6
8. Convolvulaceae 5
9. Euphorbiaceae 5 65%
10. Chenopodiaceae 4
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Importance of weeds

The most importhant weed familyes of Hungary

Family Number of species


1. Compositae 29
27%
2. Gramineae 24

3. Cruciferae 18

4. Caryophyllaceae 16 73%

5. Fabaceae 13

6. Labiatae 12

7. Chenopodiaceae 11

8. Scrophulariaceae 11

9. Polygonaceae 8 73%

10. Amaranthaceae 4

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The worst world weeds
Common name Latin name
1. Purple nutsedge Cyperus rotundus
2. Bermuda grass Cynodon dactylon
3. Cockspur grass Echinochloa crus-galli
4. Swampgrass Echinochloa colonum
5. Wiregrass Eleusine indica
6. Johnson grass Sorghum halepense
7. Red baron Imperata cylindrica
8. Waterhyacinth Eichornia crassipes
9. Purslane Portulaca oleraceae
10. Fat-hen Chenopodium album
11. Hairy fingergrass Digitaria sanguinalis
12. Field bindweed Convolvulus arvensis
13. Spring wild-oat Avena fatua
14. Pigweed Amaranthus chlorostachys
15. Spiny amaranth Amaranthus spinosus
16. Yellow nutsedge Cyperus esculentus
17. Hilograss Paspalum conjugatum
18. Itchgrass Rottboellia exaltata
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The worst weeds in Hungary
Common name Latin name
1. Cockspur grass Echinochloa crus-galli
2. Field bindweed Convolvulus arvensis
3. Fat-hen Chenopodium album
4. Yellow bristle-grass Setaria glauca
5. Redroot Pigweed Amaranthus retroflexus
6. Creeping thistle Cirsium arvense
7. Black bindweed Bilderdykia convolvulus
8. Common ragweed Ambrosia elatior
9. Field woundwort Stachys annua
10. European dewberry Rubus caesius
11. Common couch-grass Elymus repens
12. Flower of-an-hour Hibiscus trionum
13. Cornflower Centaurea cyanus
14. Green bristle-grass Setaria viridis
15. Field horsetail Equisetum arvense
16. Pigweed Amaranthus chlorostachys
17. Pale persicaria Polygonum lapathifolium
18. Charlock Sinapis arvensis
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Classification of weeds

1. Origin of weeds:
- Native
- Adventives

2. Habitats of weeds:
- Weeds adapted to breeding,
Living around the people ,
Weeds of crop production.

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Harmful aspects of weeds (Ujvrosi, 1973)

Taking of the site


Exhaustion of water store of the soil
Exhaustion of nutrient store of the soil
Decrease of soil temperature
Suppression of crop plants
Parasitic weeds

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Harmful aspects of weeds (Ujvrosi, 1973)

Weeds can harbour diseases and pathogenic fungi


Weeds can propagate pests
Weeds increase of benefits
Weeds destroy quality of crop
Poisonous weeds
Quarantine and dangerous weeds
Yield losses approach 20% caused by weeds
internationally

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Adaptation of weeds

1. Ggrowth and competition

Big photosynthetic productivity.


The most important 10 weeds of the World have C4
type photosynthesis. It means, that at first step of
photosynthesis origins an organic molecule with 4 C
atom (PEP).
Opposite to C3 type plants, which make 3-phospho-
glicerine acid in first step.

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Adaptation of weeds

1. Growth and competition

C4 plants surpass the C3 because of:


big net photosynthesis
effective water utilization (transpiration coefficient
250-350 g water /dry matter g opposite to C3 450-950
g water /dry matter g
Quick assimilate translocation
Quick growing up to generative phase.

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Adaptation of weeds

1. Ggrowth and competition


Important C4 weeds:
Amaranthus retroflexus
Echinochloa crus-galli
Setaria spp.(foxtail)
Digitaria spp.
Panicum spp.
Cynodon dactylon
Portulaca oleracea
Cenchrus pauciflorus
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Adaptation of weeds

1. Ggrowth and competition

-Weeds have rapid seedlings growth and the ability to reproduce when
young. (E.g. redroot pigweed can flower and produce seed when less
than 8 inches (3.2 cm) tall. Few crops can do this.

- Weeds have quick maturation or only a short time in the vegetative


stage. Canada thistle can produce mature seed 2 weeks after flowering.

- Weeds may have dual modes of reproduction. Most weeds are


angiosperms and reproduce by seed. Many also reproduce vegetatively.

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Adaptation of weeds

1. Ggrowth and competition

Competition for:
-Nutrients
-Light
-Water
Results in lower yields and poor crop quality

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Adaptation of weeds

2. Reproduction properties
- Weeds have environmental plasticity. Many weeds are
capable of tolerating and growing under a wide range of
climatic and edaphic conditions.

-Weeds are often self-compatible, but self pollination is not


obligatory.

- If a weed is cross-pollinated, this is accomplished by no


specialized flower visitors or by wind.
-

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Adaptation of weeds
2. Reproduction properties

- Weeds resist detrimental environmental factors. Most crop


seeds rot, if they do not germinate shortly after planting.
Weed seeds resist decay for long periods in soli and remain
dormant.

- Weed seeds exhibit several kinds of dormancy or dispersal in


time, to escape the rigors of the environment, and germinate
than conditions are most favorable for survival. Many weeds
have no special environmental requirements for germination.

- Weeds often produce seed the same size and shape as crop
seed, making physical separation difficult and facilitating
spread by man. (e.g. alfalfa and Cuscuta spp.)

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Adaptation of weeds

2. Reproduction properties

- Some annual weeds produce more than one seed crop


per year, and seed is produced for as long as growing
conditions permit.
-Each generation is capable of producing large numbers
of seeds per plant, and some seed is produced over a
wide range of environmental conditions.
-

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Adaptation of weeds
2. Reproduction properties

Roots of some weeds are able to penetrate and emerge


from deep in soil. While most roots are in the upper
foot of soil (e.g. canada tistle roots routinely
penetrate 3 to 6 feet and field bindweed roots have
been recorded over 10 feet deep. Roots and rhizomes
are capable of growing many feet per year.

- Roots and other vegetative organs of perennials are


vigorous, with large food reserves, enabling them
withstand environmental stress and intensive
cultivation.

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Adaptation of the weeds

2. Reproduction properties
Build up of soil seed bank

Seed bank can increase rapidly in one season with


fertilization and irrigation and ineffective control
One weed plant can produce thousands of seeds
Affected by management

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Adaptation of weeds
3. Allelopathic effect on crop plant

Black walnut and tomatoes


Lambs quarters roots secret oxalic acid
Velvetleaf, quackgrass, Canada thistle, giant foxtail,
black mustard and yellow nutsedge
Mechanism: root secretion, decomposition of
residues, effects microbial symbionts

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Adaptation of weeds
4. Weeds can harbour diseases and pathogenic fungi,
esp. crop relatives

Classic case of wheat rust disease on wheat with


alternative host European barberry
1970-1990 caused $100 million annually
Barberry eradication project saved farmers $30
million per year

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Adaptation of weeds
5. Can be directly parasitic
Dodder (Cuscuta spp.) Convolulaceae (Morningglory)
Family

Major problem in West US with alfalfa, clover, potatoes,


sunflower
First germinates root then when finds host becomes
parasite
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Adaptation of weeds
5. Can be directly parasitic

Striga is serious problem of corn and sorghum in Africa


Striga, commonly known as witchweed, is a genus of 28
species of parasitic plants that occur naturally in parts of
Africa, Asia, and Australia.
Indicate soil characteristics and suitability
The genus is for
classified
crops in the family
Orobanchaceae although
older classifications place it
in the Scropulariaceae.

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Adaptation of weeds
6. Indicate soil characteristics and suitability for crops:

Creeping buttercup, Ranunculus repens,


Ranunculaceae Fam.
Introduced from Europe as ornamental
Reproduces from seed or rhizomes
Toxic to cattle
Could indicate moist soil conditions

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Benefits of weeds

Enhance soil structure and water penetration


Improve soil tilts
Capture nutrients that would otherwise be lost
Provide habitat for beneficial insects
Save the soil from erosion and deflation

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Plant life-forms
The most widely applied life-form scheme is the Raunkir
system.
Raunkir plantlife-form: Danish Botanical Society in 1904 -
The subdivisions are based on the location of the plant's
growth-point (bud) during seasons

1. Phanerophytes; 2-3. Chamaephytes; 4. Hemicryptophytes; 5-9. Cryptophytes: 5-6. Geophytes; 7.Helophytes; 8-9.HydrophytesTherophites, Epiphytes

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Plant life-forms

Plant life forms by Raunkiaer

1. Woody plants (Phanerophyta) Ph


2. Dwarf shrub (Chamaephyta) Ch.
3. Partly hidden wintering (Hemicriptophyta) H.
4. Hidden wintering (Kriptophyta) K.
Wintering in soil (Geophyta) G.
Wintering in water or marsh (Hydatophelophyta) HH.
5. Annual (Therophyta) Th.

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Plant life-forms of weeds
Korsmo, E. (1930): Unkrauter im Ackerbau der Neuzeit

Balzs-Ujvrosi:
I. THEROPHYTA, T living for less than13 months annual
II. HEMITHEROPHYTA, HT living for two years biennial
III. HEMIKRYPTOPHYTA, H overwintering in the
ground level perennial
IV. GEOPHYTA, G overwintering under the ground

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Weed classification by Ujvrosi (1973)
I. One time blossoming:
- annuals:
T1 germination in autumn, seed
ripening in spring
T2 germination in autumn, seed
ripening at the beginning of summer
T3 germination in spring, seed ripening at the
beginning of summer
T4 germination in spring, seed ripening at the
end of summer

- biannuals:HT wintering with seed or rosette


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Weed classification by Ujvrosi (1973)

II. More times blossoming:


perennials:
- wintering with aboveground and
underground shoots (Ph, Ch)
- in winter the aboveground shoots are
died (H, G)

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Weed classification by life cycle
Weeds are classified into three categories based on
their life history:

Annuals complete their life cycle within one year and


reproduce from seed.
Winter annuals germinate in the fall and complete their
reproductive cycle in the spring or early summer.

Summer annuals germinate in the spring and set seed


in late summer or fall. Summer annuals thrive when
summer annual crops like corn or soybeans are
grown.

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Weed classification by life cycle

Biennials live during two growing seasons. The first


year consists of vegetative growth, while the second
year involves both vegetative and reproductive
growth. Biennials also reproduce from seed. Because
these weeds require two years to complete their life
cycles, they are found in areas of low soil disturbance
such as waterways, pastures, alfalfa, and fence rows.

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Weed classification by life cycle

Perennial plants live for more than two years.


Reproduction can occur by seed production or
vegetatively by structures such as rhizomes, tubers,
bulbs, or budding roots. Although perennial weeds
are most prevalent in areas of reduced soil
disturbance, some are well adapted to row crops.
Managing perennial weeds is generally more difficult
because of their multiple reproductive systems.

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Life cycle of an annual weed

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Winter annual (T-1,2)

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Summer annual (T-3,4)

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Biannual (HT)

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Perennials

H life form:
Overwintering organs stand vertically in the soil layer
H1: plants with fibrous root-system
H2 sarmentose plants
H3: these plants roots are able to reproduce
H4: tap-root unable to reproduce
H5: sloped root-stock

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Perennials (H)
Plalife-forms
HEMIKRYPTOPHYTA, H

Overwintering organ:

Short vertical stem on the ground Short as lope stem below ground

Fibrous root-system Taproot

Runners Reproduction organ Not reproduction organ

H1 H2 H3 H4 H5
Caltha palustris Poa trivialis Symphytum officinale Ononis spinosa Plantago major
Ranunculus acer Ranunculus repens Taraxacum officinale Eryngium campestre Artemisia vulgaris
Potentilla sp. Rumex obtusifolius Ballota nigra
Glechoma hederacea

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Perennials (H)
HEMIKRYPTOPHYTA, H1

Caltha palustris

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Perennials (H)
HEMIKRYPTOPHYTA, H2

Ranunculus repens

Glechoma hederacea

Potentilla reptans
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Perennials (H)
HEMIKRYPTOPHYTA, H3

Taraxacum officinale

Symphytum officinale
Rumex obstusifolius

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Perennials (H)
HEMIKRYPTOPHYTA, H4

Ononis spinosa
Eryngium campestre

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Perennials (H)
HEMIKRYPTOPHYTA, H5

Plantago major

Artemisia vulgaris

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Perennials

G life form:
Overwintering and reproductive organs are in the soil

G1 : Modified underground shoots stolon or


rhizome
G2: modified stem to storage, tuber
G3: propagating roots
G4: bulb or tuber-bulb

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Perennial (H, G)

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Perennials (G)
GEOPHYTA, G

Overwintering organ:

Long horizontal stem below ground Short vertical stem below ground

Stolons and rhizomes Tubers Creeping roots Bulbs

G1 G2 G3 G4
Stolons: Mentha arvensis Cirsium arvense Colchicum autumnale
Elymus repens Mentha aquatica Convolvulus arvensis Ornitogallum umbellatum
Cynodon dactylon Stachys palustris Asclepias syriaca Poa bulbosa
Calistegia sepium
Rhisomes:
Sorgum halepense
Phragmites australis

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Perennials (G)
GEOPHYTA, G1 with stolons

Elymus repens

Cynodon dactylon

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Perennials (G)
Plant life-forms
GEOPHYTA, G1 with rhizomes

Phragmites australis

Sorgum halepense

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Perennials (G)
GEOPHYTA, G2 with tubers Stachys palustris

Mentha arvensis

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Perennials (G)
GEOPHYTA, G3 with creeping roots

Convolvulus arvensis

Cirsium arvense
Asclepias syriaca

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Perennials (G)
GEOPHYTA, G4 with bulbs

Colchicum autumnale Poa bulbosa

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Weed survey methods in Hungary
Exact method's

Measuring method: the plants are cutting from a


concrete area and measured by species. It is called
for phytomass method, but it is correct when the
whole plants are measured together with roots.

Plant counting: plants are counted by species from a


concrete area.

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Estimating methods

Braun - Blanquet

Hult - Sernander

Balzs Ujvrosi

Photography from earth or air

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Braun Blanquet-scale

Resultant of dominance values dont make a straight


line (mathematically not correct).

Scale values:
1 value = less than 1/20 abundance,
2 value = 1/20 - abundance,
3 value = 1/4 - 1/2 abundance,
4 value = 1/2 - 3/4 abundance,
5 value = 3/4 - 4/4 abundance.

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Hult Sernander-scale

Values of the scale:


1 value = less than 1/16 abundance,
2 value = 1/16 - 1/8 abundance,
3 value = 1/8 - 1/4 abundance,
4 value = 1/4 1/2 abundance,
5 value = more than 1/2 abundance.

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Balzs Ujvrosi weed survey method

The method easy to study and simply to make survey,


but needs practice
We estimate the abundance of different weed
species, and finally summed abundance of sampling
area
Mathematically correct and data's are computerized
Accepted sampling area is 2x2 m2
Advantage of this method that we can check the
data's on the spot
Values can directly convert to abundance percent.

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Values of the Balzs-Ujvrosi-scale

Values Density % Values Density % Values density %

6 100.00% of survey area 3-4-4 21.87% of survey area 1-2 4.68% of survey area

5-6-6 87.50% of survey area 3-4 18.75% of survey area 1-1-2 3.90% of survey area

5-6 75.00% of survey area 3-3-4 15.62% of survey area 1 3.12% of survey area

5-5-6 62.50% of survey area 3 12.50% of survey area +-1-1 2.49% of survey area

5 50.00% of survey area 2-3-3 10.93% of survey area +-1 1.87% of survey area

4-5-5 43.75% of survey area 2-3 9.37% of survey area +-+-1 1.24% of survey area

4-5 37.50% of survey area 2-2-3 7.81% of survey area + 0.62% of survey area

4-4-5 31.25% of survey area 2 6.25% of survey area 0-+ 0.36% of survey area

4 25.00% of survey area 1-2-2 5.46% of survey area 0 0.10% of survey area

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Balzs Ujvrosi weed survey method
The weed surveying frame

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Weed survey methods
Early-season weed scouting
The first key to weed management is proper weed
identification. The best method for timely
identification is through field scouting.
The first reports on weed conditions in a field are
needed within two weeks after crop emergence to
evaluate herbicide performance and to determine if
there is a need for rotary hoeing, cultivation, or post
emergence herbicides.
Earlier scouting will be needed in no-till fields where
a knockdown or early preplant herbicide may be
applied.

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Weed survey methods

Identify and record all weed species found.


Determine the severity of the infestation by counting
the number of weeds.
Sample areas get an accurate count of the different
weeds present in the field or on the farm.
Along with weed reports, early soil moisture
observations are important. They serve as indicators
of herbicide effectiveness. Adequate moisture is
necessary for effective weed control with all soil-
applied herbicides.

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Weed survey methods

Post-emergence herbicides usually are most effective


when weeds are young and actively growing.
The degree of control with these herbicides will vary
due to differences in weed species, growth stages,
weather conditions, and herbicide application
method.
To select the best possible herbicide and apply it at
the optimum time to maximize control, the manager
needs to be able to identify weed seedlings when
they are small.

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Weed survey methods
Economic threshold for weeds
An economic threshold for weeds is the density of a
weed population at which control is economically
justified because of the potential for yield reduction,
quality loss, harvesting difficulties, or other problems
that weeds may cause.

Broadleaf and grass weeds will compete at different


levels of intensity depending upon the
competitiveness of the crop, the tillage system,
environmental conditions, and other weeds present.
In general, broadleaf weeds are more damaging to a
broadleaf crop, while grass weeds are more
competitive in a grass crop.
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Reproduction biology of weeds

Reproduction Biology of annual weeds


(By: Magyar L., Kazinczi G. 2002
Kazinczi G., Magyar L. 2003)

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Reproduction Biology of annual weeds
1. Definition of annual weeds
(Latin: annus, "year")
Annual weeds are plants that complete their life cycle [from
seed to seed] within one growing season, or in less than 12
months
survive the unfavourable season in the form of seeds
according the life-form scheme - annual species are Therophytes [Th]

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Reproduction Biology of annual weeds
Annual weed species [Therophytes] can be
classified by their life cycle as follows:
1. Winter annual species emerge in late summer or fall, survive winter, and produce
seed during late spring or early summer of the following year

2. Summer annual species emerge in spring or early summer and produce seeds
during the same growing season

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Reproduction Biology of annual weeds
2. Definition and roles of seeds
SEED:
- fertilized, mature (ripened) ovule having an embryonic plant
- it stores food material (endosperm or perisperm), and a protective coat
(testa) or coats

Roles of seeds:
1. vehicle for species multiplication
2. dispersal role
3. protection during conditions unfavourable for
germination and development (dormancy; seed bank)
4. temporary source of food for the embryo
5. source for transfer of a new genetic combinations
(diversity)

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Reproduction Biology of annual weeds
Structure of a seed

A typical seed includes three basic parts: (I.) an embryo, (II.) a supply of nutrients
for the embryo, and (III.) a seed coat
- The embryo is an immature plant from which a new plant will grow under
proper conditions. The embryo has one cotyledon or seed leaf in monocotyledons,
two cotyledons in almost all dicotyledons
- The radicle is the embryonic root
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Reproduction Biology of annual weeds
3. Seed production of weeds
extremely fluctuates (e.g Papaver rhoeas: 4-400 capsule/plant)

can be significantly different between and within species

Seed production of weeds depends on:

genotype
environmental factors and their interactions
time of germination
intra- and interspecific competition
pests, hosts, herbicides
under stress situation the life cycle can shorten

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Reproduction Biology of annual weeds
Examples of seed production by common weed species
(Stevens 1932, 1957 )

Scientific Name Number of Seeds


(per plant)

Amaranthus retroflexus 117 400


Chenopodium album 72 450
Portulaca oleracea 52 300
Capsella-bursa pastoris 38 500
Echinochloa crus-galli 7 160
Abuthilon theophrasti 4 300
Ambrosia artemisiifolia 3 380
Sinapis arvensis 1 700
Stellaria media 600

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Reproduction Biology of annual weeds
4. Dispersal of weed seeds
- Seed dispersal is the movement or transport of seeds away from the parent plant
- Most seed are good travellers, use various forces and agents to transport and
scatter themselves from place to place

What are the purposes of dispersal?

To reduce competition between the parent plant and the seeds

To avoid predators and pathogens

To provide gene flow

To maintain biodiversity

It may allow plants to colonize new habitats and geographic regions

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Reproduction Biology of annual weeds

Methods and types of seed dispersal

1. Natural: 2. Artificial:

Gravity Machinery

Ballistic (Self-dispersal) Crop seed

Water Livestock

Animal

Wind

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Reproduction Biology of annual weeds
Examples of natural seed dispersal
Gravity - Most seed of our common crop field weed species have no
specialized structures and mechanism

Water - Seed can be dispersed by water in different ways (float, flooding,


movement with surface water - irrigation, rivers, lakes)

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Reproduction Biology of annual weeds
Examples of natural seed dispersal
Ballistic (self-dispersal) - the physical and often explosive discharge of
seeds from the fruit

Wind - wings or plumes (pappus) that slow rate of fall

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Reproduction Biology of annual weeds
Examples of natural seed dispersal
Animals:
A seed eaten by an animal can experience different fates. It can be eaten and
dispersed as a viable seed, or be eaten and destroyed by digestion
a./ External way - on the body of animals (barbs, hooks, hairs)

b./ Internal way - via ingestion by animals (birds, mammals)

c./ With the help of ants - elaiosome (food body) to attract ants

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Reproduction Biology of annual weeds
Examples of artificial seed dispersal
- Human activities, most important transport mechanism is modern
agriculture
- Contaminated crop seed historically has been one of the most important
dispersial agents for weed seeds

Machinery - on equipment

Crop seeds

Livestock - feed, hay, straw and manure

Moving soil for construction and landscaping


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Reproduction Biology of annual weeds
How far can weed seeds travel?
(Mohler, 2001)

Agricultural weed seeds can travel over range of distances, depending on the method of
transport and the weed species

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Reproduction Biology of annual weeds
5. Seed Dormancy
Dispersal phase usually dormant; dehydrated seeds weight less
(esp. wind dispersal) and are metabolically slower

Definitions:

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Reproduction Biology of annual weeds

Definitions:

Dormancy can be called as "dispersal in time"

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Reproduction Biology of annual weeds

Significance:

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Reproduction Biology of annual weeds

Definitios and terms associated with seed dormancy:

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Reproduction Biology of annual weeds

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Reproduction Biology of annual weeds
The dormancy continuum
Dormancy is not a simple off-on switch

Diagram of annual changes in dormancy states of seeds


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Reproduction Biology of annual weeds
The dormancy continuum
Level of dormancy changes over time:
- seeds of summer annuals:
are dormant at maturity, and dormancy is released in spring due to low
temperatures during winter, whereas high temperatures during summer
induce secondary dormancy.

- winter annual species: require higher summer temperatures to release


dormancy for germination in autumn

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Reproduction Biology of annual weeds
Seasonality of emergence in weed species
Due to dormancy cycling most weed species germinate at particular times of year
In general, two-peak emergence differing according to species can be observed

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Reproduction Biology of annual weeds
Classification of Seed Dormancy
(Baskin - Baskin, 2004)

1. Physiological dormancy
2. Morphological dormancy
3. Morphophysiological dormancy
4. Physical dormancy
5. Combinational dormancy

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Reproduction Biology of annual weeds
1. Physiological dormancy
CAUSE physiological inhibiting mechanism of germination of embryo
prevents germination until a chemical change (hormones, phytocrome,
growth inhibitors) takes place in the seed
- divided into 3 levels - (D, I, DN)

METHOD TO OVERCOME THIS FORM OF DORMANCY cold (0-10 Co) and


warm (>15 Co) stratification

EXAMPLE most arable weed species belong to this form of dormancy


(AMARE, CAPBP, ECHCG)

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Reproduction Biology of annual weeds
2. Morphological dormancy
CAUSE - the seed is immature (underdeveloped) when shed, and a period of
growth and/or differentiation is required before germination can take place

METHOD TO OVERCOME THIS FORM OF DORMANCY provides favourable


conditions for embryo growth and germination

EXAMPLE Conium maculatum (CONMA)

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Reproduction Biology of annual weeds
3. Morphophysiological dormancy
CAUSE Seeds have embryos that are underdeveloped and differentiated
as well as a physiologocal component to their dormancy
METHOD TO OVERCOME THIS FORM OF DORMANCY these seeds
require time for embryo growth and a dormancy breaking treatment (cold
or warm stratification)
EXAMPLE some species from the families of Apiaceae, Ranunculaceae,
Papaveraceae (PAPRH)

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Reproduction Biology of annual weeds
4. Physical dormancy
CAUSE - The seed have impermeable testas or pericarps, the embryo is
therefore dry until the seed coat is broken and water enters
- seed coat may be impermeable to water, oxygen and mechanically resistant

METHOD TO OVERCOME THIS OF FORM DORMANCY chemical or physical


scarification (abrasion)

EXAMPLE commonly in hard seeds species from the families Fabaceae,


Malvaceae, Chenopodiaceae and Liliaceae
(CONAR, HIBTR, ABUTH)

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Reproduction Biology of annual weeds
5. Combinational dormancy
CAUSE - multiple mechanism, seed have simultaneously physiological (non-deep)
and physical dormancy

METHOD TO OVERCOME THIS OF FORM DORMANCY a cold or warm


stratification treatment of seeds after scarification to permit imbibition

EXAMPLE Trifolium repens (TRIRE)

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Reproduction Biology of annual weeds
Regulation of seed dormancy

Genotype + Environment = Seed Dormancy


(embryo and mother plant) (macro- and micro-)

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Reproduction Biology of annual weeds
Regulation of seed dormancy
I. Physiological regulation:

II. Role of environmental factors:


- Temperature (diurnal and seasonal fluctuation)
- Light (quality, photoperiod)
- Water availability (moisture)
- Gases (O2, CO2)
- Mineral nutrition (nitrate, NO3)
- Time of ripening and seed position

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Reproduction Biology of annual weeds
Relationship between dormancy and
phytohormones

Abscisic acid (ABA) induces and maintains seed dormancy,


inhibits seed germination

Gibberellins (GA) promote seed germination

Cytokinins, Auxins implicated in promoting seed


germination

Physiological dormancy is controlled by the ABA : GA ratio

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Reproduction Biology of annual weeds
Photoregulation of seed dormancy
The seeds can detect: - absence or presence
- quality (wavelength composition)
- intensity
- duration of light

Positive, negative photoblastic and light insensitive species

Photoreceptor in seeds = PHYTOCHROME SYSTEM


- 5 different Phy (A-E) types of different physiological function

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Reproduction Biology of annual weeds
Action mode of Phytochrome
The phytochrome system is the photoreceptor of red light responses
PR - physiologically inactive form - it absorbs red (R; 670 nm) light;
PFR- physiologically active form - it absorbs far red (FR; 730 nm) light

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Reproduction Biology of annual weeds
What is a weed seed bank?
Mature seeds are shed from the parent plant and find themselves on the soil surface
The term seed bank is used to describe the
reservoir of viable seeds present on the
surface and in the soil
It consists of numerous seeds many of which
are dormant and delay germination until a
latter time
The size of a seed bank in agricultural land
varies enormously (between sites, fields and
plots) from near zero to as much as 1 million
seeds per m2
It includes
New seeds recently shed by plants
Older seeds that have persisted in the soil
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Reproduction Biology of annual weeds
Importance of weed seed banks
It is an indicator of past and future weed problems
It is the primary source of new infestations of weeds each year
It gives help to
- learn the dynamics of weed population
- predict plant invasion
- plan the methods and strategies of weed management

Only 1-9 % of the viable seeds produced in a given year develop into
seedlings, the rest remain viable and will germinate in subsequent
years depending on the depth of their burial

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Reproduction Biology of annual weeds
Composition of seed banks

A few species dominate the seed bank


They are primary weeds in a cropping system
Adapted to cropping system
Resistant to control measures
Many infrequent species
Adapted to your area but not to current production
practices
Include
Newly introduced species
Species survived previous land uses

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Reproduction Biology of annual weeds
Types of seed banks
(Thompson and Fenner, 2005)

On the basis of the length of time that seeds survive in the soil a seed bank
can be:
Transient - seeds persist in the soil for less than 1 year
e.g. GALAP, BROMO, CENCY
Short-term persistent - seeds persist in the soil at least
1 year but less than 5 years
e.g. TAROF, CONCA
Long-term persistent - seeds persist in the soil more
than 5 years
e.g. STEME, CAPBP, CONAR, PAPRH, AMARE
_________
Seed size + morphological feature The type of a seed bank
- small (0,5-1 mm) + flat surfaced seeds = persistent seed bank
- large (2-4 mm) + appendage seeds = transient seed bank
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Reproduction Biology of annual weeds
What happens to seeds in the soil?
Active versus dormant (persistent) seed banks
Active seed bank consists of seeds that are able to
germinate
Dormant seeds make up the persistent seed bank and
are able to survive for many years
Examples:
Small seeded annual weed seeds without a hard seed coat only
persist a couple of years e.g. SET sp., DIGSA

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Reproduction Biology of annual weeds
Dynamics of soil seed bank

(inputs to seed bank are shown with black arrows, and losses in white arrows)

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Reproduction Biology of annual weeds
Weed seed rain
Weed seed input (also called seed rain)
Without seed input the seed bank will rapidly
decline
Local sources of seed predominates
95% of seed rain comes from annual weeds growing in
field
Distant sources are important if local seed production
is limited

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Reproduction Biology of annual weeds
Seed persistence
The longevity of seeds in the soil is highly variable, and depends on species,
depth of seed burial, soil type and the level of disturbance
most long-lived species are annuals
small seeds tend to have much longer soil lives than large ones
max. seed viability in the soil e.g. Spergula arvensis: 1600 years (in archaeological digs)

Weed species survival (years)


______________________________________________

Cirsium arvense 21
Chenopodium album 39
Portulaca oleracea 30
Ambrosia artemisiifolia 39
Amaranthus retroflexus 10
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Reproduction Biology of annual weeds
Weed seed distribution in the soil
Majority of seeds in no-till agricultural
field is located in the upper 5 cm of
soil profile
In cultivated soils in the upper 15 cm
of soil profile
Cultivation and tillage continuously
invert the soil profile
Bringing up seeds previously buried to
the upper soil surface layers more
readily germinate
The type of cultivation is important for
distribution (e.g. ploughing buries
seeds below 10 cm)

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Reproduction Biology of annual weeds
Tillage effects on weed seed distribution
Repeated tillage for several years without reinfestation
will reduce the weed seed population in soil

However, if weeds produce many seeds, then it is


extremely difficult to completely avoid reinfestation

Conclusion: allowing weeds to have seeds increases


potential problems for many years to come

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Reproduction biology of weeds

Reproduction Biology of perennial


weeds

1
Reproduction Biology of perennial weeds
(Latin per, "through", annus, "year")
A perennial plant or perennial is a plant that lives for
more than two years
Perennials are Herbaceous and Woody perennials
(trees and shrubs)

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Perennial weeds

Herbaceous perennials grow and bloom over the


spring and summer and then die back every autumn
and winter, then return in the spring from their root-
stock rather than seeding themselves as an annual
plant does

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Perennial weeds

Many invasive weeds


are perennials

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Reproduction of perennials

Reproduction of perennials:

I. Strategies of reproduction
II. Main asexual forms of reproduction
III. Factors affecting the regeneration of perennials

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Reproduction of perennials
I. Strategies of reproduction
a) Mainly reproducing by seeds
High seed production and effective seed propagation.
Spreading by seeds but settlement by asexual ways.
Rumex crispus, R. obtusifolius, Ranunculus repens, Mentha arvensis,
Achillea millefolium
b) Generative and vegetative forms are equal
importance
Tough, hard to kill weeds.
Have to control both the seedling and the daughter
plant. Sorghum halepense, Asclepias syriaca, Calystegia sepium

c) Mainly reproducing vegetatively


Weeds of no-or low tillage, or wrong cultivation
Stolons can grow 9,9 m a year (ELYRE)
Elymus repens, Cynodon dactylon, Cirsium arvense

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Reproduction of perennials
II. Main asexual forms of reproduction

a) Stolons and rhizomes

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Reproduction of perennials
II. Main asexual forms of reproduction

Stolon

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Reproduction of perennials (Elymus repens)
II. Main asexual forms of reproduction
Stolon

(Hunyadi et al.1988)

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Reproduction of perennials
II. Main asexual forms of reproduction
Stolon

Principles of stem growth of common reed (Haslam, 1972)

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Reproduction of perennials
II. Main asexual forms of reproduction
Rhizomes

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Reproduction of perennials
II. Main asexual forms of reproduction
Rhizomes

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Reproduction of perennials
II. Main asexual forms of reproduction
Rhizomes

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Reproduction of perennials
II. Main asexual forms of reproduction
b) Creeping roots

The reproduction organ never


goes above ground
(like rhizomes)

Root system of Convolvulus arvensis


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Reproduction of perennials
II. Main asexual forms of reproduction
b) Creeping roots

Root system of Cirsium arvense

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Reproduction of perennials
(

Root system of Cirsium arvense

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Reproduction of perennials
II. Main asexual forms of reproduction
c) Reproducing from taproot

Rootcrown

Axillarybuds Sideshoot
leaf

Adventitiousroots

Taproot
Lateralroot

Modified taproot of Rumex obtusifolius


(Hunyadi et al. 1988)
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Reproduction of perennials
II. Main asexual forms of reproduction
c) Reproducing from taproot

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Reproduction of perennials
II. Main asexual forms of reproduction
d) Tubers, bulbs, runners

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Reproduction of perennials

Tubers and bulbs

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Reproduction of perennials
II. Main asexual forms of reproduction

Importance of asexual reproduction:

More nutrients for progeny

Exact reproduction of the parent genotype

Small a mount of progeny

No ability to long-distance spread

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Reproduction of perennials

III. Factors affecting the regeneration of perennials

1. Periodicity

2. Apical dominance

3. Environmental effects

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Reproduction of perennials

III. Factors affecting the regeneration of perennials

1. Periodicity

Periodicity is a seasonal activity of buds on the vegetative organ


First observation on Agropyron repens stolons
Late Spring Dormancy
(Johnson and Buchholtz, 1957, 1958, 1962,
Confirmed by Hakkanson, 1967; Leakayet al., 1972 and Hunyadi,
1978)
Seasonal activity also descibed on Cynodon dactylon (Hunyadi),
Sorghum halepense(Hunyadi) and Phragmites australis (Lukcs-
Hunyadi)
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Reproduction of perennials
III. Factors affecting the regeneration of perennials
1. Periodicity

Seasonal activity of buds on 1-node AGRRE stolon segments

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Reproduction of perennials
III. Factors affecting the regeneration of perennials
1. Periodicity

Seasonal activity of budson1-node CYNDA rhizome segments

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Reproduction of perennials
III. Factors affecting the regeneration of perennials
1. Periodicity

Seasonal activity of budson1-node PHRAU rhizome segments

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Reproduction of perennials
III. Factors affecting the regeneration of perennials
1. Periodicity

Seasonal activity of buds on1-node SORHA rhizome segments

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Reproduction of perennials

III. Factors affecting the regeneration of perennials


2. Apical dominance

Inhibition of one bud to an other

On an uncultivated area 90% of the axilary buds are


dormant

The axilary buds are inhibited by the apex


Apical Dominance

Buds closer to the apical part developing longer shoots

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Reproduction of perennials
III. Factors affecting the regeneration of perennials
2. Apical dominance

Regeneration of a 7-nodes AGRRE stolon after segmentation

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Reproduction of perennials
III. Factors affecting the regeneration of perennials
2. Apical dominance

Importance:

Apical Dominance is a preventing-mechanism of a perennial


plant after segmentation by cultivation or after bud death
caused by herbicide

the plant can recover easily and quickly

Keeping most of the buds dormant, perennials can save


lots of stored nutrients by controling the flow

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Reproduction of perennials

III. Factors affecting the regeneration of perennials


3. Environmental effects

Depth of a perennating organ

Temperature

Light

Nitrogen

Size of the perennating organ


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Effects of environmental factors on
weeds
Factors relating to weed establishment and survival

Environmental factors:
1. Climatic factors:
Light (intensity, quality, and duration including
photoperiod)
Temperature (extremes , average, frost-free period)
Water (amount, percolation, runoff and evaporation)
Wind (velocity, duration)
Atmosphere (CO2, O2, humidity)
Many of the most common weeds have a broad
tolerance to environmental conditions

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Factors relating to weed establishment and survival

2. Physiographic factors
Edaphic:
soil factors
pH
Texture
Structure
Fertility
Organic content humus
Water drainage
CO2, O2
Topographic:
altitude
slope
exposure to the sun

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Factors relating to weed establishment and survival

3. Biotic factors:
Plants:
competition
allelophaty (released toxins or stimulants)
parasitism
diseases
soil flora
Animals:
insects, grazing animals, soil fauna,
Humans

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Forms of plant interactions
1. Competition
2. Allelopathy
3. Parasitism: the phenomenon of one living organism living in,
on or with another living organism to complete its life cycle
an one-sided association in which one of the symbionts (the
parasite) benefits at the expense of the other (the host).
4. Symbiosis: symbiosis is the living together of different
organisms, usually in close association with one another, to
the benefit of at least one of them. The partners are
referred to as symbionts.

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Factors relating to weed establishment and survival

Weed populations are dynamic in nature and can


change rapidly in response to different management
practices such as herbicide application and tillage
systems. It is important to understand the underlying
mechanisms that enable different weed species to
adapt to different selection pressures.

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Competition
Competition

Weed competition is a part of weed echology.


Competition in echology involves two or more
organisms seeking for a particular factor, thing or
material when they are in short or limited supply
(Donald 1963).

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Competition
Type of competition

Interspecific between varieties


Intraspecific between individuals of variety
Intergenotypic - between genotypes of variety
Intragenotipyc between individuals of genotype
Intraindividual within one individual plant
( perennial weeds, f.e. competition for water between
sprouts of shoot and rootstalk at Asclepias syriaca. It
is an important factor to preserve the sleeping status
of the sprouts on rootstalk)

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Competition

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Competition

Weed compete with crops for environmental


resources available in limited supply:
Nutrients
Water
Light
Territory/space

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Influencing factors of competition
Competitive capacity of a varieties is genetically
determined.
Attributes influenced competition: (Clements et al.
1929) For them competition was a purely physical
process.
1. Permanency: seizing of territory and height
2. Rhythm of growth
3. Quick germination
4. Vigor favorable under stress to survive

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Influencing factors of competition

Influencing factors:
1. Reaction to different light intensity
2. Reaction to rising temperature
3. Reaction to changing oxygen of atmosphere
4. Photorespiration (presence or absence)
5. C3 or C4 photosynthesis

Black et al. 1969

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Influencing factors of competition
Shematic diagram of the competition encountered by
a plant

(Blesdale 1960)

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Influencing factors of competition
Pozsgai 1988: factors of competition must complete
with anthropogenic effect:
agrotechnic
Fertilizers
Plant protection
Weed control
Integrated plant protection give preference to crop
plant

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Critical competitive period

All crop plants can tolerate weeds for a period


The period while must provide without weeds for
crop call for critical competitive period.
It depend on species, growing technology (seed sowing
or plantation), weed flora, etc.

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Critical competitive period some crops
Critical competitive period of some crops

week
Maize 3
Rice 3
Peanut 3-4
Soya bean 3
Wheat 5
Cotton 6
Potato 9
Sugarbeet 8
Oilseed rape 10
Onion 12
Flax 12
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Competition for nutrients
Fertilization is used to improve crop growth, but may
worsen the weed problem.
Macronutrients: nitrogen, potassium, phosphorus,
primary plant nutrients
Nitrophyl weeds: Galium aparine, Apera spica-venti, Avena
fatua, Chenopodium album, Amaranthus retroflexus
Mezo- and micronutrients content of weeds usually
higher than crops

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Competition for nutrients
Kilograms of nutrients required to produce equal
amounts of dry matter

Plant Nitrogen Phosphorus


Wheat 5,5 1,2
Oats 4,9 1,7
Barley 8,4 2,6
Common lambsquarters 7,6 1,6
Common ragweed 6,6 1,4
Redroot pigweed 5,1 1,4
Common purslane 3,1 0,8
Mustards 9,8 2,7

Weeds require the same nutrients, at the same times, and they usually
take up nutrients more quickly, and greater extent than crop plant in
ecosystem.

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16
Competition for nutrients
Nitrogen is the first nutrient to become limiting in most
instances of weed-crop competition. Rooting depth and root
area of plant determine the ability to obtain resources, and
relative competitiveness for nitrogen is largely determined by
the soil volume occupied by roots of competing species.
Movement of P and K is slow, and they move over short
distance. Competition most likely to occur after plant are
mature and have extensive, overlapping root development.
Because of this fact competition for P and K more apparent in
perennial crops.
Concentration of N and K usually highest at beginning of
vegetation period, but P at generative period.

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16
Szraz hajtstmeg (g/nvny)

12
14
16
18
20

10

0
2
4
6
8
Kontroll

Stomp 330+Basagran

Pledge 50 WP+Basagran

N0
Command 48 EC+Basagran

Kontroll

Stomp 330+Basagran

Pledge 50 WP+Basagran

N100

TMOP-4.1.2.A/2-10/1-2010-0012
Command 48 EC+Basagran
Competition for nutrients

Kontroll

N200 Stomp 330+Basagran

Pledge 50 WP+Basagran

Command 48 EC+Basagran

Kontroll

Stomp 330+Basagran
N300

Pledge 50 WP+Basagran
Dry matter production of pea influenced by nitrogen and herbicides

Command 48 EC+Basagran
16
Szraz hajtstmeg (g/nvny)

10
15
20
25
30
35
40
45
50

0
5
Kontroll

Stomp 330+Basagran

Pledge 50 WP+Basagran

N0
Command 48 EC+Basagran

Kontroll

Stomp 330+Basagran

N100
Pledge 50 WP+Basagran

TMOP-4.1.2.A/2-10/1-2010-0012
Command 48 EC+Basagran
Competition for nutrients

Kontroll

Stomp 330+Basagran

N200
Pledge 50 WP+Basagran

Command 48 EC+Basagran

Kontroll

Stomp 330+Basagran
N300

Pledge 50 WP+Basagran

Command 48 EC+Basagran
Dry matter production of Chenopodium album influenced by nitrogen and herbicides

16
Hajts nitrogntartalom (%)

0
1
2
3
4
5
6
Kontroll

Stomp 330+Basagran

Pledge 50 WP+Basagran

N0
Command 48
EC+Basagran

Kontroll

Stomp 330+Basagran

N100
Pledge 50 WP+Basagran

TMOP-4.1.2.A/2-10/1-2010-0012
Command 48
EC+Basagran
Competition for nutrients

Kontroll

Stomp 330+Basagran
N200

Pledge 50 WP+Basagran

Command 48
EC+Basagran

Kontroll

Stomp 330+Basagran
N300

Pledge 50 WP+Basagran

Command 48
EC+Basagran
Nitrogen content of Chenopodium album influenced by nitrogen and herbicides

16
sszes gyomborts (%)

10
20
30
40
50
60

0
Kontroll

Command 48
EC+Basagran
Sencor 70 WG+Basagran

N0
Stomp 330+Basagran

Kontroll

Command 48
EC+Basagran

N1
Sencor 70 WG+Basagran

TMOP-4.1.2.A/2-10/1-2010-0012
Stomp 330+Basagran
Competition for nutrients

Kontroll

Command 48
EC+Basagran
N2

Sencor 70 WG+Basagran

Stomp 330+Basagran

Kontroll
Effect of nitrogen and herbicides on weed density

N3

Command 48
EC+Basagran
Sencor 70 WG+Basagran

Stomp 330+Basagran
16
Competition for water
Water or its lack is often the primary environmental
factor limiting crop production, and water is probably
the most critical of all plant growth requirements.
Without irrigation the rainfall determines the
geographic limit of crops.
Weeds compete for water, reduce water availability,
and contribute to crop water stress. Weedy
sunflowers require approximately twice as much
water as corn.

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16
Competition for water
In arid areas developed a fallow cropping system:
wheat is grown one year and the land is fallowed (no
crop) next year, and rotated back to wheat in the
third year. The aim is water conservation, because
there is not enough rainfall to growth wheat each
year, therefore minimum or no-tillage system,
sometimes called ecofallow. The ecofallow system
increased water, soil nitrate, grain protein and wheat
yield.

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16
Competition for water

Weeds have different strategies to tolerate water stress


Terminology of Levitt (1958) distinguish two form of drought
suffering:
Arid-active plants: restrict their water losses on different
ways
Arid-tolerant plants: they can lose relatively lot of water
without damage. Subletal water deficit of Ambrosia
artemisiifolia and Digitaria sanguinalis is above 70%. They
can loss 70% of their water content without irreversible
injury (Mikuls et al. 1991)
- Smaller shoot-root ratio of weeds is advantage in contrast to
crops

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17
Competition for water

Roots grow more rapidly, than shoots.


Competition greatest when roots closely intermingle
and crops and weeds try to obtain water from the
same volume of soil.
Less competition occurs if roots of crops and weeds
are concentrated in different soil areas
More competitive faster-growing plant have large
root system so they are able to exploit a large volume
of soil quickly.

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Competition for water

Some species save their water content with efficient


stoma regulation, or with leaf area reduction (leaves
of mezofil grasses roll upon spiral form, or fall down,
and later growth new leaves from latent sprouts) .

Leaves developed under stress condition have less


leaf area, have thicker cuticle, and more carved
leaves

Under water stress weeds able to shorten their


vegetative period and to ripe the seeds

C4 photosynthetic type plants use less amount of


water to make dry matter than C3 plant.
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17
Competition for light
The total supply of light is the most reliable of the
several environmental resources required for plant
growth.
But in contrast to water and nutrients, light cannot be
stored for later use- it must be used when received or
it is lost forever.
Light regulates many aspects of plant growth and
development.
It varies in duration, intensity and quality.
Neighboring plants may reduce light supply by direct
interception -shading

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17
Competition for light
Leaves that first intercept light may reflect it, absorb
it, convert it to photosynthetic products, convert it to
heat, or transmit it.
If transmitted, the light is filtered so that when is
reaches lower leaves, it is dimmer and spectrally
altered.
Any time one leaf is shaded by another, there is
competition for light.
Light competition is most severe when there is high
fertility and adequate moisture because plants grow
vigorously and have larger foliar areas.

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Competition for light
Plants with large LAI have a competitive advantage
and normally outcompete plants with smaller leaf
area.

Successfull competitors do not necessarily have more


foliage, but have their foliage in the most
advantageous position for light interception. (plant
with opposite leaves are probably less competitive
than those with alternate leaves. Plant that are tall or
erect have a competitive advantage for light over
short, prostrate plants.

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Competition for light
A heavily shaded plant suffers reduced
photosynthesis, leading to poor growth, a smaller
root system, and a reduced capacity for water and
nutrient uptake.

Crops and weeds differ in shade tolerance. Yellow


nutsedge grown between corn rows than within the
row because less light reaches the soil under plants,
so yellow nutsedge density decreases as corn density
increasesmanagement technique is increasing plant
population, but it is limited!

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17
Plant characteristics and competitiveness
Foliar competition
Rapid expansion of a tall, foliar canopy
Horizontal leaves
Large leaves
C4 photosynthesis
Leaves forming a mosaic leaf arrangement for best
light interception
A climbing habit (Gallium aparine, Convolvulus
arvensis)
Tall stem
Rapid stem extension in response to shading

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17
Plant characteristics and competitiveness
Root competition:
early and fast root penetration of a large soil area
High root density/soil volume
High root-shoot ratio
High root length per root weight
Actively growing roots
Long and abundant root hairs
High uptake potential for nutrients and water

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17
Effect of weed competition on yield

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17
Effect of weed competition on yield

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18
Methods to study competition
1. Replacement designs
Total plant density is maintained constant, but the relative
proportion of species is varied from zero to a pure stand.
Additionally, for each species, pure stands must be included at
each of densities in the mixtures to enable intraspecific
competition to be determined.
This method often undertaken in glasshouse pot studies.

plant number in pots


crop weed
4 0
3 1
2 2
1 3
0 4

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18
Methods to study competition

2. Additive designs
To study the effect of increasing weed density on crop yield we use
most often the additive design. We can predict of yield loss.
Density of one species (usually the crop) is maintained constant while
the other (weed) varies.
This method we use in glasshouse pot studies and on field experiments.

plant number
crop weed
4 0
4 1
4 2
4 3
4 4

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18
Methods to study competition

TMOP-4.1.2.A/2-10/1-2010-0012

18
Leaf area of pea and Sinapis arvensis in an additive experiment

600
A bors levlfellete A SINAR levlfellete

500
levlfellet (cm2/edny)

400

300

200

100

0
Kontroll SINAR 5 db/t.e. SINAR 10 db/t.e. SINAR 15 db/t.e.

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18
Methods to study competition
3. Growth analysis

From the point of view theory and practice equally


very important to know the growth stages of plants.
BASF, Bayer, Ciba-Geigy and Hoechst companies
made a decimal scale to code the development of
monocotyledonous and dicotyledonous plant, calls
BBCH scale
We study increase of dry mass and leaf area in a
determined phase.

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18
BBCH scale
Subdivision of the developmental cycle of plant

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18
Methods to study competition

3. Growth analysis

Increase biomass
production

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18
Methods to study competition

Relative Growth Rate, RGR

Relative Leaf Growth Rate, RLGR

Net Assimilation Rate, NAR

Leaf Area Ratio, LAR

Crop Growth Rate, CGR

Specific Leaf Area, SLA

Leaf Weight Ratio, LWR

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18
Allelopathy
Allelopathy
The word allelopathy:
Derived from Greek allelo, meaning each other and patho,
an expression of sufferance or disease.

Molisch defined the term of allelopathy in 1937.


Allelopathy refers to chemical interaction between
plants (microbes and higher plants), including
stimulatory as well as inhibitory influences (Rice 1974).
Rice (1984) called interference every interaction between
plants.
Allelopathy is a part of interference
allelopathy means secretion of allelochemicals by donor plant,
and these allelochemicals stimulate or inhibit acceptor plant.

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19
Allelopathy

Allelopathy is a kind of stress.


In the plant community, plants exert adverse or
depressive effect on their neighbors by releasing toxic
substances into the immediate environment.
Allelopathy is believed to be a significant factor in
maintaining the present balance among the various
plant species.

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19
Allelopathy part of competition

We know for a long time, that there are plant species


in natural succession, which hinder the colonization
of other species.
In this case increase of biomass production we can
see only on one species, so we cant explain this
symptom with competition for nutrients or water.
This is allelopathy.

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19
Early observations

Theophrastus (BC. 300): weed suppressive effect of pea


Demokritos (BC. 400.): suggests to use plant material
as a weed managing method
Plinius Secundus (A.D. I.C.): injuring shadow of walnut
for plants and humans
smell and pressed liquor of some plants dangerous for
radish, and laurel harms grape-vine

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19
Allelopathy

Allelopathy very important in tropical areas, because


of accumulation of plant residues in the soil, and
because have no enough time for organic matter
decomposition.
Allelopathy research: USA, India
Themes:
Study of natural ecosystems
Weed-crop interaction
Identification of allelochemicals

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19
Chemistry of allelochemicals

Plants produce numerous metabolites of no known


utility to their growth and development.
These are often referred to as secondary plant
metabolites, and are often defined as compounds
having no known essential physiological function.
Allelochemicals vary from simple molecules such as
ammonia to the more complex organic molecules.

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19
Chemistry of allelochemicals
Allelochemicals in plants

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19
Juglon index:
Juglone: powerful toxin produce by the leaves of black
walnut (Juglans nigra) 5-hydroxy--naphthaquinone
hinders emergence

Juglon index: (Szab 1999)


Compares the effect of plant with unknown
allelopathy to 1 mM juglon treatment. Test species is
white mustard (Sinapis alba L.)
Quotent of germination percentages, or root and
shoot lengths determine the juglon
>1 allelopathic potencial stronger, than juglons
< 1allelopathic potencial poorer, than juglo

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19
Production of allelochemicals

They are produced by any plant organ:


Roots
Leaves
Seeds
Flowers
fruits

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19
Production of allelochemicals
Allelochemical compounds are released from the
plants:
Vapor
Leaching from the foliage
Exudates from the roots
Decomposition of dead plant residues

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19
How allelochemicals get out of plant

(Aldrich, 1984)
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20
How allelochemicals get out of plant
Evaporation the most important in arid and semiarid
areas of the world. (Artemisia spp., Eucalyptus spp.)
Exudates from the roots:
Verified with wheat, oak, maize, pea, cucumber, tomato,
sunflower
Ambrosia artemisiifolia, Sorghum halepense, Dygitaria
sanguinalis

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20
How allelochemicals get out of plant
Leaching from the foliage:
Pinus densiflora, Juglans regia, Helianthus annuus,
Platanus occidentalis, Brassica nigra, Abutilon theophrasti

Decomposition of dead plant residues: -by microorganisms


Very hard to decide that inhibiting compounds leaching from
the plant residues or originated from transforming effect of
microorganisms

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20
Weeds with allelopathic activity
Weeds with alleged allelopathic activity

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20
Weeds with allelopathic activity
Weeds with alleged allelopathic activity
Weed susceptible species

TMOP-4.1.2.A/2-10/1-2010-0012

20
Weeds with alleged allelopathic activity
Weeds with alleged allelopathic activity
Weed susceptible species

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20
Crop plants with allelopathic activity

Crop with alleged allelopathic activity


Crop susceptible species

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20
Crops whose residues are phytotoxic

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20
Crops whose residues are phytotoxic
Crops with phytotoxic residues

Crop susceptible species

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20
Effect of allelochemicals
Direct:
Effect on germination, growth, metabolism
Indirect:
Effect on soil properties, nutrient content, change of plant
population , microorganism population, insect population

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20
Allelochemicals influence:
Cell division, and ultra structure
Plant hormones and their balance
Permeability of membranes
Germination of pollens and spores
Nutrient uptake
Stomach regulation, photosynthesis, pigment synthesis
Respiration
Protein synthesis
Nitrogen fixation
Enzyme activity
Tissue formation
Water circulation
Genetic program
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21
Allelochemicals
Allelochemicals from plants with potential
herbicidal activity

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21
Allelochemicals
Volatile essential oils

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21
Allelochemicals

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Allelochemicals

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21
Allelochemicals
Category of compound weeds affected

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21
Allelochemicals
Category of compound weeds affected

TMOP-4.1.2.A/2-10/1-2010-0012

21
Allelochemicals
Category of compound weeds affected

TMOP-4.1.2.A/2-10/1-2010-0012

21
Allelochemicals
Category of compound weeds affected

TMOP-4.1.2.A/2-10/1-2010-0012

21
Allelochemicals

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21
Effect of allelochemicals on ecosystems

Scientific results prove, that allelopathy has an


important roll in natural and artificial ecosystems.
- in development of plant succession (primeval
grasses, deforestation) toxins are very important:
-appear and disappear of species
- changing of species dominance
- disappear of pioneer weed species
- reforestation
- vegetation under the leafy boughs

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22
Effect of allelochemicals on ecosystems

Nitrogen fixation and nitrification hindering effect:


Successions develop in the nitrification hindering
and ammonium conservating direction in a
climax succession the population of Nitrosomonas
and Nitrobacter species are very small.
More important weeds hinder the development of
Rhisobium species tubercles on the roots of
Leghuminacea

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22
Effect of allelochemicals on ecosystems
In a lot of ecosystems live only one species, or density
of species is characteristic
Among phanerophytas walnut, locust, plane tree
have a selective herbicidal effect on weeds
Soil boreding: problem at orchard replanting
(peach, apple)
After growing of Sorghum cant growing other plants

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22
Methodologies for studying allelopathy

Making extraction from donor plant:


Cut or chop the leaves roots, etc.
Using solvents (acetone, ethanol etc.)
Or cold water
Rate of plant and solvent mass usually
4-100:100
Extraction time 24-48 hours
Filtration
Storage: some days on 4-5 C

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22
Methodologies for studying allelopathy
Bioassay

Controlled examination under laboratory conditions


Dilution row
Germination test in Petri dishes:
Percentage of germination
Length of germ
Suggested test plants: bean (Phaseolus vulgaris), Vicia faba,
rape (Brassica napus), oats, Lepidium draba

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22
Bioassay
Winter wheat

Ambrosia seed extract

Ambrosia leave extract

Control

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22
Bioassay

Radish control

Radish+ABUTH leaves
extract

Radish+ ABUTH seed


extract

TMOP-4.1.2.A/2-10/1-2010-0012

22
Bioassay

Bean control

Bean+ABUTH
leaves extract

Bean+ABUTH
seed extract

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22
Bioassay

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22
Pot experiments in green house
Use different soil types
1. Extracts may use one time (unit area)
- or watering more times
2. mixed the dried leaves or roots into the
soil
Estimation:
we measure the germination rate, hight of plants,
fresh- and dry veight, nutrient content, etc.

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22
Step methode

Weed
Crop

(Radosevich Holt, 1984)

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23
Field experiments
On small plots
Environmental factors have an important roll: temperatur,
rain, etc.
Microbiall degradation
Spraying solution
Or sowing donor and acceptor species together
Mixed plant parts into the soil, or took to the soil
surface

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23
Methode to determine allelochemicals

(Tang Young, 1982)

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23
Cultural practices:
1. Crop rotation and weed management
The manipulation of allelopathic rotation crops can
provide an effective means of weed management.
- sunflower reduces the density of both broadleaf and
grassy weeds.
- soybean-corn-wheat rotation decrease of giant
foxtail population
- alfalfa reduces the incidence of weeds in the next
crop (Avena fatua, cirsium arvense, Gallium aparine)

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23
Cultural practices:
2. Allelopathic cover crops
Cover crops are grown for several motives such as
conservation of soil and moisture, improvement of
nutrient cycling, lowering of temperatur, supplying
forages in emergency, protection of cash crops from
winds, suppression of weeds, increase in crop
production.
Hairy vetch suppresses weeds, reduce the emergence of
chenopodium album, imperata cylindrica.
Velvetbean and kudzu can effectively control speargrass
(Imperata spp.) in fallows and maiz fields.

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23
Cultural practices:
Cover crops with allelopathic property and weed-
supressing ability

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23
Cultural practices:

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Allelochemicals identified from the cover crops

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23
Cover crop allelochemicals

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23
Cultural practises:
Green manure crops

Legumes- soybean, clovers, Mucuna spp.)


Cruciferous plants (Brassica spp. mustards). Leaves
from rapeseed incorporated into soil reduce
broadleaf weeds

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23
3. Allelopathic crop residues
The presence of crop residues on the soil surface as
mulch suppresses weeds through allelopathy
- Wheat (Amaranthus spp.) aqueous extracts inhibit
the growth of velvetleaf, barnyardgrass, and redroot
pigweed
- Rice: grassy and broadleaf weeds
- Sorghum: Digitaria ishaemum, velvetleaf, redroot
pigweed and Rumex acetosella
- Alfalfa: grassy and broadleaf weeds

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24
Allelochemicals as herbicides
Allelopathy has the potential to play a prominent role
in weed management if it can be properly harnessed
Characteristics:
They are environmentally friendly
They have comparatively less residual activity, rapid
degradation
They have novel target sites of action

Allelochemicals can be used in either crude or pure


form.

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24
Allelochemicals as herbicides
In spite of many compounds being tested, only a few
have been marketed.
mesotrion leptospermone

Callistemon citrinus

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Herbicide resistance in weed species
Definition of resistance

Weed resistance is the inherited ability of a plant biotype to


survive the application of a herbicide, which had originaly
been effective against that weed population.

Susceptible weeds must be distinguished from tolerant ones.

The susceptible weed is a weed biotype that is unable to


survive the application of a herbicide at its normal rate.

The tolerant or unsusceptible weed is a biotype that has


never been affected by the use of a herbicide.

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Definition of resistance
Cross resistance:
resistance to more than one chemicals with the same target
site or same metabolic enzime.
For example, one herbicide selected for the resistant biotype, but this
biotype is also resistant to other herbicides in the same mode of
action.

Multiple resistance:
occurs when resistance to several herbicides results from
two or more distinct resistant mechanism in the same
plant.
For example, resistance to both triazine and ALS-inhibiting herbicides,
which are two different modes of action.

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Definition of resistance

Herbicide resistant plants can often resist high doses of a herbicide.


This shattercane biotype is not affected by a dose 64X that
recommended for Beacon (primisulfuron; ALS-inhibitor).

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Definition of resistance

Weed species such as common cocklebur resistant to one


herbicide (eg. Classic, chlorimuron) may also be resistant to
another herbicide (e.g. Scepter, imazaquin) in the same mode
of action (ALS-inhibitor). This is known as cross-resistance.

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Definition of resistance

Common waterhemp seedlings were identified as having


multiple resistance, that is, resistance to both triazine and
ALS-inhibiting herbicides, which are two different modes
of action with different mechanisms conferring resistance.

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Distribution of herbicide resistant weeds

The first case of herbicide resistance (to 2,4-D) was reported by Hilton (1957).
The first case of atrazin resistance in Senecio vulgaris (to simazine) in Washington
State was reported by Ryan (1970).

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Distribution of herbicide resistant weeds

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Distribution of herbicide resistant weeds

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Herbicide resistant weeds

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Distribution of herbicide resistant weeds

Distribution of cases of herbicide resistance in developing and developed countries according to


MoA (B.E. Valverde: Herbicide-resistance management in developing countries)

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Distribution of herbicide resistant weeds

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Development and mechanism of resistance in weeds

Development and mechanism of resistance in


weeds

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Development of resistance

In most of the cases resistance evolve by the


continued use of a particular herbicide

The development of resistance is not correlated to


plant families or genus

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Development of resistance
Important factors help to evolve herbicide-resistance:

1. The frequency of the resistance genes


The genes conferring resistance are present naturally in wild
populations at a very low frequency. This frequency is
important in determining how long it would take for the
resistance to become noticable.

2. The selection pressure imposed by the herbicide


Higher selection pressure is imposed when we use herbicides at
high doses and we spray them too frequently. The efficacy,
persistency, specificity of a herbicide are also affecting
factors.
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Development of resistance

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Development of resistance
Important factors help to evolve herbicide-resistance:
3. The weed fitness
Ecological fitness is the ability of one biotype to be more successful in a given
environment than another because of greater biomass production, more
seed produced per plant, or higher germination potential. This has been
observed for res. biotypes of weeds to triazine herbicides. If no herbicide is
applied and res. and selective biotypes are competing, the proportion of
resistant biotype should decrease since they have reduced ecological
fitness

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Development of resistance

4. The size and the viability of the soil seed bank


The soil seed bank may act as a buffer, thus delaying
the evolution of resistance. The importance of this
natural or less selected genetic seed bank is
depending on the lifetime of the seeds, the
germination characteristics of the weeds and the
cultivation technology of the crop.

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Mechanism of resistance

1.Target site insensitivity

2. Enhanced herbicide metabolism or breakdown


to inactive products

3. Herbicide sequestration

4. Overproduction of the target site

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Mechanism of resistance

1. Target site insensitivity

There are specific target sites for herbicides. The


structure of these enzymes can be modified by a
mutation. The herbicides can not bind to the site
anymore. Cross resistance often occur.

Examples: resistance to triazines, ALS-inhibitors,


dinitro-anilines

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Mechanism of resistance

1. Target site insensitivity

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Mechanism of resistance

1. Target site insensitivity

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Mechanism of resistance

2. Enhanced herbicide metabolism

Three groups of enzymes have been implicated in


metabolism-based resistance:
glutathione transferases (GSTs)
aryl acylamidases
cytochrome P450 mono-oxygenases

Resistance to atrazine in Abutilon theophrasti

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Mechanism of resistance
2. Enhanced herbicide metabolism
Enzyme System Weed Species Herbicide
Cytochrome P450 monooxygenase Alopecurus myosuroides chlorotoluron
diclofop
propaquizafop
chlorsulfuron
Avena sterilis diclofop
Lolium rigidum diclofop
chlorsulfuron
chlorotoluron
metribuzin
simazine
chlorotoluron
Phalaris minor isoproturon
Sinapsis arvensis ethametsulfuron
Stellar media mecoprop
Glutathione transferases Abutilon theophrasti atrazine
Alopecuris mysouroides fenoxaprop
Aryl acylamidase Echinochloa colona propanil
Echinochloa crus-galli propanil

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Mechanism of resistance
3. Herbicide sequestration

Physical or temporal separation of the herbicide from sensitive


tissues or target sites.

Example: Crassocephalum crepidioides found in Malaysia,


resistant to paraquat

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Mechanism of resistance
4. Overproduction of the target site

This mechanism is identified in tissue culture selections, as the


resistance mechanism to graminicides (ACCase inhibitors) in a
johnsongrass (Sorghum halepense) biotype.

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Resistance by mode of action

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Resistance to triazines

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Resistance to triazines

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Resistance to triazines
Triazines are inhibiting photosynthesis at PSII by binding to QB (D1)
and blocking the transport of electrons between Q-PQ.
Resistance is based on several mechanism, mainly:
Modified binding site of QB-protein (target site insensitivity)
Hydroxilation, dealkylation, glutation-conjugation (herbicide
metabolism)
First ocurence: Amaranthus hybridus (USA Maryland) 1972

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Resistance to triazines
QB (D1) -protein
PQ (Plastoquinon)

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Resistance to triazines
QB (D1)-protein

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Resistance to triazines

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Resistance to ALS-inhibitors

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Resistance to ALS-inhibitors
Three groups are inhibiting the acetolactate-synthetase:
imidazolinones (IMI), pyrimidinylthiobenzoates (PTB),
sulfonylaminocarbonyltriazolinones (SCT), sulfonylureas (SU),
and triazolopyrimidines (TP).

These herbicides are inhibiting protein synthesis by inhibiting


the synthesis of 3 branched-chain amino acids Valin, Leucin
and Isoleucin. The first enzyme of this pathway is ALS.

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Resistance to ALS-inhibitors
Multiple mutations are responsible for ALS resistance. For
example, mutations in the gene that encodes ALS which result
in any one of the amino acids changes will result in resistance
to the ALS inhibitors. The particular mutation determines to
which ALS inhibitor family the weed will be resistant.
For example: change in Pro197 sulfonylureas,
substitution at Ala122 imidazolinone
Plants resistant to the ALS inhibitors do not have reduced
fitness. Therefore, the resistant populations build up quickly.

First ocurence: Lactuca seriola (USA - Idaho) 1987

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Resistance to ALS-inhibitors

The gene responsible for resistance can be transplanted to


crop plants.

For example:
Resistance to sulfonil-ureas SUMO and Expres varieties
Resistance to imidazolinones - IMI varieties

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Resistance to glyphosate

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Resistance to glyphosate

Glyphosate kills plants by inhibiting EPSP-Synthase in


meristematic tissue. It is a key enzyme in the
shikimate biosynthetic pathway which is necessary
for the production of the aromatic amino acids
(fenilalanine, tirozin, triptofan), auxin, phytoalexins,
folic acid, lignin, plastoquinones and many other
secondary products.

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Resistance to glyphosate

Glyphosate occupies the binding site on EPSPS for


phosphoenol pyruvate, a substrate of EPSPS, by
mimicking an intermediate state of the enzyme-
substrates complex.

The two mechanisms of resistance are


1) alterations of the target site, EPSPS, and
2) decreased uptake/translocation of
glyphosate
to the meristematic tissues.

First occurence: Lolium rigidum (Australia Victoria) -


1996
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Resistance to glyphosate

There are two forms of EPSPS in nature, EPSPS I, which


is found in plants, fungi, and most bacteria, and is
sensitive to glyphosate, and EPSP II, which is found in
glyphosate resistant bacteria and is not inhibited by
glyphosate. It is the gene for an EPSPS II that has
been used to genetically engineer resistance in crops.

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Resistance to ACCase-inhibitors

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Resistance to ACCase-inhibitors
The enzyme Acetyl-coenzyme-A-carboxylase catalyses the first
committed step of fatty acid biosynthesis.

This enzyme is the target site of two groups:


aryloxyphenoxypropionates (AOPP) and cyclohexanediones (CHD).
These herbicides kill monocot weeds but do not affect dicot
weeds.

Resistance is conferred by an ACCase alteration that results in


reduced herbicide sensitivity.
There can be resistance due to rapid metabolism in some species.

First occurence: Alopecurus myosuroides (UK) - 1982

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Management of herbicide
resistant weeds

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Management of herbicide resistant weeds

The prevention of resistance occurring is an easier and


cheaper option than managing a confirmed resistance
situation.

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Management of herbicide resistant weeds
The following key areas of weed management will help to reduce the
selection pressure on any weed species hence significantly reducing
the chance of survival of resistant weeds:

1. Cultural techniques

Cultivation or ploughing prior to sowing to control emerged plants


and to bury non-germinated seed
Delaying planting so that initial weed flushes can be controlled with a
non-selective herbicide
Using certified crop seed free of weed
Post harvest grazing, where practical
Stubble burning, where allowed, can limit weed seed fertility
In extreme cases of confirmed resistance, fields can be cut for hay or
silage to prevent weed seed set

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Management of herbicide resistant weeds

2. Crop rotation

Different crops will allow rotation of herbicides having a different


mode of action

The growth season of the weed can be avoided or disrupted

Crops with differing sowing times and different seedbed preparation


can lead to a variety of cultural techniques being employed to
manage a particular weed problem.

Crops also differ in their inherent competitiveness against weeds. A


strongly competitive crop will have a better chance to restrict
weed seed production.

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Management of herbicide resistant weeds
3. Herbicide rotation and herbicide mixtures

Avoid continued use of the same herbicide or herbicides having the


same mode of action in the same field unless it is integrated
with other weed control practices

Limit the number of applications of a single herbicide or herbicides


having the same mode of action in a single growing season

Where possible, use mixtures or sequential treatments of herbicides


having a different mode of action but which are active on the
same target weeds

Use non-selective herbicides to control early flushes of weeds (prior


to crop emergence) and/or weed escapes

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Management of herbicide resistant weeds

3. Herbicide rotation and herbicide mixtures

For chemical mixtures to be effective, they should:

Include active ingredients which both give high levels of control


of the target weed, AND

include active ingredients from different mode of action


groupings.

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Management of herbicide resistant weeds

Additional to the above guideline, the grower should


Know which weeds infest his field or non-crop area and where
possible, tailor his weed control program according to
weed densities and/or economic thresholds
Follow label instructions carefully. This especially includes
recommended use rates and application timing for the
weeds to be controlled
Routinely monitor results of herbicide applications, being
aware of any trends or changes in the weed populations
present
Maintain detailed field records so that cropping and herbicide
history is known

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Management of herbicide resistant weeds
What to do in cases of confirmed herbicide resistance
The degree of the action will depend on the stage of the crop in the field and
the extent of the problem.

Eradicate the remaining weed population if growing in patches in order to


limit build-up and spread of seed in the soil
Limit the field to field movement of resistant populations by cleaning
planting, cultivation and harvesting equipment to avoid transfer of resistant
weed seed
Avoid using the herbicide to which resistance has been confirmed unless
used in conjunction with herbicides having a different mode of action, active
on the resistant weed population
If the resistant population is widespread consider grazing the crop or cut for
feed being careful not to transfer resistant seed via manure
Select these fields for rotation or set aside for the following cropping
season

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Useful books and sites:
R. Zimdahl: Fundamentals of weed science
R. Zimdahl: Weed-crop competition: a rewiew
T. A. Hill: The biology of weeds
F. A. Macias, Juan C.G. Galindo, Jose M. G. Molinillo, Horace G. Cutler:
Allelopathy: Chemistry and Mode of Action of Allelochemicals
CRC Press September 2003
Wod Powel Anderson: Weed Science, principles and applications
A.H. Cobb J.P.H. Reade: Herbicides and Plant Physiology
Robert E. L. Naylor (ed.): Weed Management Handbook
Prado et al.: Weed and Crop Resistance to Herbicides
R. Labrada (ed): Weed Management for Developing Countries
http://www.fao.org/docrep/006/y5031e/y5031e00.htm#Contentsc
Plant and Soil Sciences eLibrary
http://plantandsoil.unl.edu
Plant Physiology online
http://5e.plantphys.net/
The HRAC website
http://hracglobal.com/
International survey of herbicide resistant weeds
http://www.weedscience.org
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Francisco A Macias, Juan C.G. Galindo, Jose M. G. Molinillo, Horace G Cutler 2003:
Allelopathy: Chemistry and Mode of Action of Allelochemicals
CRC Press
Magyar L., Kazinczi G. 2002: A gyommagvak nyugalmi llapota s
csrzskolgija. I. A magnyugalmi llapot (dormancia) okai, tpusai s
feloldsnak lehetsgei. Magyar Gyomkutats s Technolgia 3 (2), 3-20.
Kazinczi G., Magyar L. 2003: A gyommagvak nyugalmi llapota s
csrzskolgija. II. A gyommagvak szntfldi csrzsa s a dormancia fajon
belli eltrse. Magyar Gyomkutats s Technolgia 4 (1), 3-17.

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Thank you for your attention
Dr. Ndasyn dr. Ihrosi Erzsbet
Dr. Rita Szab
Georgikon Kar
Nvnyvdelmi Intzet

AZ ELADS LETLTHET:
-

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