Allelopathy

Quintin R. Johnson October 13, 1999 University of Delaware Research & Education Center Georgetown, DE

History
Theophrastus (ca. 300BC ) chick pea exhausts the ground. Molish (1937) - coined term allelopathy
biochemical interactions between all types of plants including microorganisms includes both inhibitory and stimulatory effects
most compounds inhibitory at some concentrations are stimulatory to the same processes in very small concentrations.

Terminology
Allelopathy - in current use generally restricted to detrimental effects. Allelochemicals - toxic substances released by some plants into the immediate environment of other plants that harm or kill them. May be secondary metabolites.
Phytoinhibitins - higher plant origin Saproinhibitins - microbial origin

Terminology
Interference - general term - the effect that the presence of a plant has on the growth or development of its neighbors. Negative Interference
Competition - mutually adverse effects of organisms (plants) that utilize a resource in short supply. (- , -)

Terminology
Negative Interference (cont.)
Amensalism - A dramatic decrease in plant biomass or mortality evident for one species but not for another - So striking that it cant be explained by resource competition alone. (0 , -)
Includes inhibitory affects of allelopathy. Allelopathy is distinguished from other forms of interference in that the effect is exerted through the release of a chemical by the donor plant

Terminology
Parasitism - one plant lives in or on another and derives resources directly from its host. (+ , -)

Autotoxicity - allelochemicals produced by a plant that are toxic to itself.

classic example - alfalfa, forage grasses

.Heterotoxicity - allelochemicals produced by a plant that are toxic to other plants.

Allelopathy vs. other forms of Interference

Difficult to separate from other forms of interference - especially resource competition Difficult to prove direct effects of allelochemicals vs. indirect influence via other organisms or microenvironmental modifications lack of techniques in field studies.

Proving Allelopathy Kochs Postulates Approach

Demonstrate existence of Interference
describe symptoms and quantitate degree

Isolate, assay, characterize, and synthesize toxins (allelochemicals). Characterize their biological activity. Repeat symptoms of interference with toxins applied at rates naturally present and timings appropriate to observed interference.

Proving Allelopathy
Monitor release, movement, and uptake of toxins - Show sufficient concentrations for observed interference. Beware of the possibility of synergistic effects of more than one toxin. Consider species selectivity of toxins if possible. Proof vs. strong supporting evidence?
Proof not yet obtained in the field

Sources of Allelochemicals
Volatilization
Primarily associated with arid region plants Artemisia, Eucalyptus, Salvia absorbed in vapor, from dew condensate, or by roots after reaching the soil

Root Exudation
occurs in many plants active exudation, leakage, sloughed dead cells? Absorbed by plant roots

Sources of Allelochemicals
Leaching
from aerial plant portions by rainwater or fogdrip chrysanthemum, flaxweed, velvetleaf understory in forest situations

Decomposition of Plant Residues

direct release from plant tissue that has lost cell integrity microbial enzymatic degradation

Sources of Allelochemicals
Decomposition of Plant Residues
Important in modern agronomic systems utilizing cover crop and/or crop residue management (i.e.. minimum and no-tillage systems)
maintenance of high levels of plant residues affects crop emergence, growth, and productivity affects weed germination, growth and competitiveness

Allelopathic Agents
Toxic Gasses (cyanide, NH3, ethylene) Organic acids and Aldehydes (citric acid, acetic acid) Aromatic Acids (cinnamic and vanillic acid) Simple Unsaturated Lactones (Patulin, parasorbic acid) Coumarins (coumarin, esculin, psoralen) Quinones (juglone)

Allelopathic Agents
Flavonoids (phlorizin, diosmetin trioside) Tannins (gallic, ellagic and digallic acids) Alkaloids (cocaine, caffeine, chinin, codein) Terpenoids (camphor, cineole) and Steroids (digitoxigenin, strophanthidin) Others?

Production of Allelochemicals Environmental Conditions

Light quality, intensity, and duration
IR light and UV light increase production of some compounds
supplemental UV light in greenhouse studies?

Red and far red light? Longer days may increase production Day and nighttime temperatures?

Production of Allelochemicals Environmental Conditions

Mineral Deficiencies
Nutrient deficiencies may enhance or limit allelochemical production. Boron, Calcium, Magnesium, Nitrogen, Phosphorus, Potassium, Sulfur Nitrogen & Phosphorus
can cause large increases - may be important in revegetation of infertile fields. Excess phosphorus?

Production of Allelochemicals Environmental Conditions

Water Stress
little work has been done generally cause increase in production

Temperature
warmer temperatures may increase production chilling may also increase production

Pesticide applications (herbicides, growth regulators) - Additive effect? Soil pH?

Production of Allelochemicals Environmental Conditions

Age of plants and growth stage Genetics
breeding programs and or bioengineering?

Pathogens and predators Combinations of stress factors

antagonism or synergism on production?
Extent and Duration Dependent on compound, species, stress

Manipulation through stress management?

Allelochemicals Mode of Action

Alter nutrient uptake
may be pH dependent

Inhibit cell division Inhibit cell enlargement Inhibit photosynthesis Stimulate or inhibit respiration Alter membrane permeability or integrity Inhibit enzyme activity

Allelochemicals Fate in Soils

Immobilization in soil organic matter
biologically active?

Accumulation on soil colloids

Higher accumulations in heavier textured soils Increased duration of activity in heavier textured soils

Chemical and/or microbial decomposition

Chemical (non-microbial) decomposition? Microbial decomposition - fungi, bacteria

Allelochemicals Effect on Ecosystems

Influence on Plant succession
germination inhibition seedling toxicity

Inhibition of nitrogen fixation and nitrification - selection toward plants that inhibit nitrification - climax vegetation patterning (spatial distribution) - zones of inhibition - selection towards pure populations or stands

Allelochemicals Effect on Ecosystems

Prevention of seed decay and germination
dormancy characteristics resistance to microbial decay Weed seed bank characteristics?

Replant problems in cropping systems

(e.g. apple orchards, continuous sorghum)

Allelopathy and Weed Management

Superior weed suppressing crop varieties
germplasm screening conventional breeding or genetic transfer cucumber, oat, sunflower, soybean high density monoculture crops (I.e turfgrass, forage grasses, legumes)

Rotational and cover or companion crops

suppression from living mulches, residues

Herbicide discovery

Techniques Monocultures
Density dependent phytotoxicity
with finite toxin availability, lower densities have more toxin available per plant over time than at higher densities greater biomass yields at higher plant densities than at lower densities = evidence of toxicity possible deviation from law of constant total yield at high densities when resource competition occurs.

Techniques Mixed Cultures

Replacement Series
Density-dependent effects on competition ignored

Target-neighbor design
vary densities of neighbor species planted around target plant in presence of a toxin. greater biomass yields at higher neighbor densities than at lower densities = evidence of toxicity

Techniques Root Exudates

Stair-step System
Donor and Recipient plants grown in separate pots in sand solution with pots alternated in stairstep fashion water, nutrients, light, and other growth factors can not be limiting Donor and recipient in alternating column compared to Donor only and recipient only columns

Weeds from our list with shown allelopathic potential

Smooth Pigweed Common Ragweed C. Lambsquarters Morningglory Sp. C. Cocklebur Velvetleaf Jimsonweed E. B. Nightshade Giant Ragweed Penn. Smartweed Burcucumber Spurred Anoda Prickly Sida Horseweed C.E. Primrose Wild Mustard Field Pansy

Weeds from our list with shown allelopathic potential

Smooth Pigweed Common Ragweed C. Lambsquarters Morningglory Sp. C. Cocklebur Velvetleaf Jimsonweed E. B. Nightshade Giant Ragweed Penn. Smartweed Burcucumber Spurred Anoda Prickly Sida Horseweed C.E. Primrose Wild Mustard Field Pansy

Weeds from our list with shown allelopathic potential

Foxtail Species Fall Panicum Large Crabgrass Johnsongrass Goosegrass Barnyard Grass Texas Panicum Bermudagrass Yellow Nutsedge Canada Thistle Horsenettle Common Pokeweed Common Milkweed Hemp Dogbane Curly Dock Trumpet creeper Field Bindweed Bigroot Morningglory

Weeds from our list with shown allelopathic potential

Foxtail Species Fall Panicum Large Crabgrass Johnsongrass Goosegrass Barnyard Grass Texas Panicum Bermudagrass Yellow Nutsedge Canada Thistle Horsenettle Common Pokeweed Common Milkweed Hemp Dogbane Curly Dock Trumpet creeper Field Bindweed Bigroot Morningglory