Weed Biology
JA Dille, Kansas State University, Manhattan, KS, USA
Ó 2017 Elsevier Ltd. All rights reserved.
This article is a revision of the previous edition article by R.E.L. Naylor, volume 3, pp. 1485–1494, Ó 2003, Elsevier Ltd.
Introduction
In this article, weeds are defined and the common features of
their biology described. Population dynamics of weeds is
considered and a brief outline of options for managing weeds
is given.
What Is a Weed?
A simple definition of a weed is a plant growing where it is not
wanted. As humans, we have decided which plants are desir-
able in a given location, and which plants are detrimental to
our desired goal. In a crop production situation, the presence
of weeds can have a harmful effect on the desired crop’s yield
or quality. In a flower bed or vegetable production situation,
presence of weeds can detract from the esthetic value, make it
difficult to harvest the vegetables, and reduce the quality of
the produce. The Weed Science Society of America defined
a weed as “a plant growing where it is not desired.” The Euro-
pean Weed Research Society uses the following definition: “a
weed is any plant or vegetation, excluding fungi, interfering
with the objectives or requirements of people.”
A weed is often a plant out of place, such that a corn plant
growing in the field where it was planted is desirable, but when
it appears in a soybean crop the following year, it is considered
a weed. Thus, any plant species and any type of plant can be
considered a weed. It is very difficult to describe all traits of
all plants that might be considered weedy.
Weedy plants can be found in very different environments
across different taxonomic classifications, including aquatic,
terrestrial, and aerial environments. In aquatic environments,
blue-green algae and macrophytes such as elodea (Elodea cana-
densis) can be troublesome plants. In aerial environments,
bromeliads such as airplants (Tillandsia) take advantage of
stressed trees and open leaf canopies to grow on tree branches.
Most weedy plants occur in terrestrial environments such as
summer annual broadleaf and grass weeds in crop fields, peren-
nial and brushy weeds in range and pasture lands, dandelion or
mosses in urban lawns or sports turf, mosses or liverworts in
ornamental nursery pots and creeping perennials such as field
bindweed or quackgrass in gardens. Taxonomically, plants
considered weeds could be algae, mosses and liverworts, ferns
and horsetails, to gymnosperms (eastern red cedar or salt
cedar), or angiosperms (flowering plants).
What is the biology of weeds? By knowing unique charac-
teristics for an individual plant species, we can understand
where will it grow well based on a location’s environmental
conditions of temperature, moisture, and day length; we can
use information about its lifecycle (i.e., annual, biennial, or
perennial) to know when it will occur in a growing season
and how long it will survive; does it reproduce by seed or
vegetatively or both, and how does it spread from location
to location.
Encyclopedia of Applied Plant Sciences, 2nd edition, Volume 3 http://dx.doi.org/10.1016/B978-0-12-394807-6.00026-5
What Traits Make Plants a Weed?
What Are Harmful and Beneficial Aspects of Weeds?
Inherent in the definition of a weed is the idea that it is harm-
ful, to other crops, humans, livestock, and the environment.
However, it is just a plant with unique traits, some of which
result in harmful aspects and some in beneficial aspects. If we
could determine alternative uses for the plants known as
weeds, they would not be classified as such and could provide
benefits. Losses due to weeds can be categorized as direct and
indirect losses. Direct losses would include those that reduce
both the quantity and quality of the ‘crop’ being produced.
These losses are associated with weed presence in any agricul-
tural production situation, such as row crops, vegetable and
fruit crops, ornamental and trees, turf and athletic fields,
pasture and rangeland, or natural environments. Indirect losses
would include those that do not reduce cash return from the
crop being produced, but represent a cost to society at large
or to the producer or property owner. For example, harboring
insects, diseases and pests of other crops, creation of health
hazards such as pollen from common ragweed that results in
hay fever suffering in humans or dermatitis by poison ivy
leaf toxins; reduced property values due to weed invasion,
and cost of weed maintenance in noncrop environments,
rights-of-way, public lands, and so on.
The main measure of harm is in lost yields. Increasing weed
densities in row crops cause increasingly greater losses of
yields. Even with adequate weed control efforts, weed presence
can still cause significant loss (10–25%) of grain yields. The
next main harmful measure is reduced quality of harvested
product that could be due to contamination by weed
seed (jointed goatgrass in winter wheat), weedy plant parts
(Canada thistle heads in canned garden peas), seed staining
(eastern black nightshade in soybean). Often this contamina-
tion results in extra costs for cleaning the product or dockage
for the crop when sold. Presence of weeds can result in harvest-
ing difficulties, such as remaining green while the crop senes-
ces prior to harvest and not able to be cut and processed
through a combine harvester or other such machine. If weeds
are present during hand-harvesting of vegetable and fruit
crops, it interferes with efficiency. Additional costs include
processing of harvested crop and/or cost of transporting
weed seed with grain, or loss of storage efficiency with green
plant parts in harvested crop.
Weeds are known to harbor insect and disease pests as alter-
native hosts and as a ‘green bridge’ between different crops in
a rotation sequence to increase pest management costs,
possible yield losses, in a subsequent crop. For example,
numerous soybean pests use weeds as alternate hosts, such as
Asian soybean rust on kudzu, soybean aphids on buckthorn,
and soybean cyst nematodes on winter annual weeds such as
purple deadnettle and henbit. Presence of green foxtail
provides a bridge between winter wheat crops by harboring
aphid pests that transfer viruses.
469
470 Weeds and Competition j Weed Biology
There are a few weedy plants that are parasitic on beneficial
plants and can result in reduced crop growth and yield reduc-
tions, such as field dodder (Cuscuta spp.) in alfalfa or clover
crops; broomrape (Orobanche spp.) which attacks carrots,
tomatoes, sunflowers, redclover, and witchweed (Striga spp.
mainly S. hermonthica) that can infect corn and sorghum and
drastically reduce their grain yields by 10% compared to
a healthy crop. Dwarf mistletoe (Arceuthobium spp.) is a parasite
on pines and firs while broadleaf mistletoe ( Phoradendron mac-
rophyllum) infects numerous landscape trees; each reduces
wood quality, increases chance for disease infection, and results
in death of the trees.
Presence of weeds in aquatic environments can prevent
water flow by blocking ditches and irrigation canals, and tran-
spire water that could have been delivered to crop plants. The
water hyacinth ( Eichhornia crassipes) is an important tropical
aquatic weed. Eurasian watermilfoil ( Myriophyllum spicatum)
is considered to be originally from Europe and Asia, released
into North American lakes and rivers and has drastically altered
the ecology of infested water bodies. The sheer mass of plants
can cause flooding, and the stagnant mats are good habitat
for mosquitoes. Milfoil mats can rob oxygen from the water
by preventing the wind from mixing the oxygenated surface
waters to deeper water. The dense mats of vegetation can also
increase sedimentation rates by trapping sediments. Milfoil
interferes with power generation and irrigation by clogging
water intakes. It interferes with recreational activities such as
swimming, fishing, and boating. The growth of some aquatic
weeds may reduce oxygen content of the water and lead to
fish deaths.
Harmful aspects of weeds on humans include discomfort
due to allergies causing ‘hay fever’ (pollen from trees, grasses,
and weeds such as common ragweed (Ambrosia artemisiifolia)
and due to dermatitis caused by the oily sap from poison ivy
(Toxicodendron radicans), poison oak (T. diversilobum), or
poison sumac (T. vernix) that contains urushiol and when on
skin, results in an itchy rash anywhere from a few hours to
days after exposure.
There are many weed species that impact animal production
and animal health. Animals can die from eating poisonous
plants, off-flavor is imparted to milk and other dairy products
when weeds such as wild garlic (Allium vineale) are eaten, and
livestock hides and carcasses are physically damaged by weeds.
Plants with spines or thorns, such as thistles, inhibit animal
foraging and protect neighboring plants from being eaten,
thus reducing the area available for grazing by livestock.
Some animals, such as horses, are much more sensitive to
certain plants than others, and if these plants are present in
degraded pasture areas so that they are grazed by horses, such
as poison hemlock (Conium maculatum), tansy ragwort (Senecio
jacobaea), yellow starthistle and Russian knapweed (Centauria
spp.), they can experience chronic effects if enough is
consumed, and result in animal death.
It may seem strange to consider the positive effects of weeds;
however, weeds are just plants and only from human perspec-
tive, are they troublesome. Beneficial aspects of weeds include
pollination and nectar resources for bees, monarch butterflies,
and other insects, source of genetic traits that could be intro-
gressed into crop plants for improved fitness, biomass produc-
tion that could be used as biofuel resources such as field
pennycress (Thlaspi arvense) for oil or switchgrass (Panicum vir-
gatum) for cellulosic biofuel, used for coloring and dyes, and
a component of biodiversity in agricultural areas. Many crop
fields in North American agriculture are monocultures of row
crops such as maize and soybean, thus described as having
reduced biodiversity. In other places, policies are in place to
encourage farmland diversity either by reducing intensity of
management or by reducing the farmed area through set aside,
such as a current European Union policy. By these practices,
weedy plants contribute to increased biodiversity.
There is interest in designing weed management systems that
could enhance the beneficial aspects of weedy plants, for
example, that would allow for development of populations of
beneficial organisms, such as pollinating insects, seed-eating
birds, insects, and mammals, and harbor natural predators,
such as ground beetles and spiders, of aphid pests of crops.
For example, if a certain level of weeds in a crop could be toler-
ated, this may allow for more farmland birds to survive and to
rear chicks before these food sources are removed. Weedy plants
in stubble fields provide a level of erosion control from wind
and water if there is little residue. However, successful weed
control practices would still need to be available prior to estab-
lishing a crop in these fields. Concerns would be raised if there
was significant presence of herbicide-resistant weed populations
allowed to grow uninhibited.
What Are the Population Dynamics of Weeds?
Weed species can be categorized into different life cycles,
including annual (winter or summer), biennial, and perennial
(simple, creeping, or woody). Important features of each life
cycle include time of seed germination, number of seedlings
established, rate of vegetative growth, time of flowering and
seed production, and seed dispersal. Annual weeds complete
their life cycle within a period of 12 months or less, while bien-
nial weed species require up to 24 months, and perennials
survive longer than 2 years, sometimes not producing flowers
and/or seed for several years until plants are well established.
Perennials often have the capability of regenerating from vege-
tative structures (roots, adventitious buds, tubers, rhizomes,
stolons, and so on) in addition to seed.
Weed Seed Dispersal in Space and Time
The biology of annual and biennial weed species emphasizes
reproduction by seed. Very high levels of seed production per
plant are common along with unique dispersal mechanisms
in space via wind, water, animals, and humans, as well as
dispersal mechanisms in time via dormancy. Seed dispersal
by plants in space is a passive process, in the sense that plants
cannot actively choose where their offspring will go. However,
various features of the seed itself and the maternal plant can aid
dispersal. Taller plants may disperse seeds over greater
distances. Lighter seeds may be dispersed further by wind,
but may have a lower chance of becoming a new seedling
because of limited stored energy reserves. Heavier seeds fall
near the maternal plant, where they may have a greater chance
of landing in a favorable site for germination since it was
a successful location for the parent, but could be detrimental

due to future competition with siblings or increased likelihood
of predation by seed predators. In general, most seed fall near
the parent plant if no active mechanism is assisting the
dispersal. To achieve greater distances, some seed or fruit con-
taining the seed can be moved internally by animals or because
of unique appendages moved externally when attached to the
animal. For the weed species, the advantage of dispersal in
space is to find new sites, while a disadvantage might be the
absence of another plant needed to achieve cross pollination
for successful seed production. The establishment of a soil
seedbank provides a means for the weed species to have
different generations and genetics that can appear in future
years (see article Weeds and Competition: Weed Seed Biology
for more details on seed rain, soil seedbank, and seed
dormancy/germination).
Modeling the Population Dynamics of Weed Species
Understanding the overall dynamics of a weed population
provides insights for making management decision based
on when the population might be most vulnerable, but not
on an expectation that the measure will be successful. Often,
the seedling growth stages are targeted for weed control early
in the cropping season, but if these seedlings only represent
a small fraction of the total soil seedbank, another aspect of
the population should also be targeted, such as the amount
of seed produced by mature plants and subsequently returned
to the seedbank each year, or by manipulating the placement
of seed in the seedbank to reduce their longevity. Many of the
models of weed populations are based on information on
reproduction of a limited number of weed species and are
used to compare the efficacy of particular management
options over a long term. The models may be particularly
useful in appraising the likely effectiveness of combinations
of management decisions (concerning use of tillage practices,
time of crop planting, selection of crop type, management of
fallow periods, and so on) in alternative scenarios, but inevi-
tably can only be as good as the data on which the models are
based. Models can be used to assess the efficacy of interven-
tion at different stages in the life cycle of weeds. They may
assist in decision making on whether to use a cheaper herbi-
cide that is less effective, rather than on a more effective but
more expensive one. Models can help in deciding whether
or not to treat the weed, that is to incorporate a weed
‘threshold’ into the management decision in order to prevent
unnecessary or uneconomic treatments.
If weeds have been managed well in an intensive agricul-
tural system, individual species often show patchy or scattered
spatial distributions among fields. Weed management by some
farmers might lead to local extinction of a given weed species
(but often difficult because of the soil seedbank and persistent
seed for some species). However, reinvasion can easily occur
due to dispersal of seed by wind, water, animals, and/or
humans. The small, sometimes transient, populations in a land-
scape might be termed a metapopulation. The dynamics of the
metapopulation (multiple small patches that are intercon-
nected via dispersal) will depend partly on the size of the local
populations or patches, and their rate of extinction. The
connectivity among patches is important and determined by
dispersal rate and distance between patches that can determine
Weeds and Competition j Weed Biology 471
the reinvasion rate. These factors can lead to large-scale
temporal and spatial variations in overall and local population
sizes.
Weed Introduction (to New Sites)
Seed dispersal by plants is a passive process, but various plant
and seed features aid in their dispersal. The goal of seed
dispersal in space is to seek out new and better environments
for future generations, to aid in genetic mixing among indi-
viduals, and to not be competing with parent plant or future
‘siblings.’ A drawback may be the absence of another related
plant with which to achieve cross pollination for production
of new seed. Individual weed species produce differing
numbers of seed, and seed weight indicates amount of stored
energy reserves. There is usually a simple trade-off between
number of seed produced and individual seed weight. Thus,
plants could produce many lightweight seed to be dispersed
widely by wind as compared to few heavyweight seed that
fall near the parent plant. Taller plants may disperse seed
over greater distances from the parent plant. Many seeds
also have adaptations for animal dispersal (internal or
external to the animal) that encourage even greater dispersal
distances. Fleshy fruits are aided by birds or other species
that eat the fruit and transport the seed long distances before
being deposited. Externally, burs and seed with sticky coats
attach to animal fur or human clothing and are transported
until rubbed off.
(For features of ‘seed rain,’ the seedbank in the soil, and the
topics of seed dormancy and germination, see Weeds and
Competition: Weed Seed Biology.)
Managing Weeds
Weed management is covered in more detail in the article. A
successful weed management strategy does exploit our knowl-
edge of weed biology to achieve crops with minimal presence
of weeds, levels known to not affect crop productivity beyond
the economic cost.
Weeds, like crops, are living organisms that respond to their
environment. We manipulate the environment to produce
conditions conducive for growth of crop plants, but weeds
also exploit these optimal conditions. The optimal plant envi-
ronment should give the advantage to the crops over the weeds.
Weed management is one of the key considerations among
many for successful crop production, and each decision and
choice made influences the likely weed species and infestation
levels that will be observed in their fields. The time of the
previous harvest, management of the residues of the previous
crop, type and timing of soil tillage, fertility management,
choice of crop species and variety, date of planting, and row
spacing and seeding rates are all decisions made about crop
production that will influence the type of weed population
that appears in the field.
An integrated approach to developing weed management
systems is recommended, and this includes knowledge of the
biology of the weed species occurring in the field, determining
the level of infestation (is management even necessary?), and
then evaluating numerous options. These control tactics can
472 Weeds and Competition j Weed Biology
be categorized as cultural, physical, biological, and chemical
options.
Assessing weed populations in the field can be problematic.
First step is correct identification of the weeds species, and this
should be done at the seedling stage of growth if timely control
has to be implemented. The distribution of weeds in a field
tends to be patchy, thus requiring the entire field to be scouted.
The implication of weed patchiness is that the application of
control can be localized rather than across the entire field. If
infestation levels are relatively low, it will be inappropriate to
determine an average infestation for a whole field that results
in underestimating weed populations in high-density patches
and recommending weed control in weed-free or low-density
areas.
Weeds in the Future
Weed species are very adaptable. Changes in crop management
and in climate are likely to alter the occurrence of weed species
in fields. Climate change may result in the shift of weed species
occurrence so that ‘new’ weeds from other areas move either
as a direct response or as a response to changes in cropping
systems. Invasions of new species into areas will generate
new challenges for growers. Our global economy often encour-
ages movement of plant species around the world, and thus
potentially introductions to new areas. Reliance on the
repeated use of single herbicides or single herbicide modes
of action has led to the evolution of herbicide-resistant weed
biotypes, which have proved challenging to manage.
Continued focus on understanding weed biology and use of
integrated weed management systems should provide effective
means of managing these plants.
See also: Arable Crops: Agricultural Crops; Agroecology:
Farming Systems with Nature as Guide. Plants and the
Environment: Invasive Plant Species. Seed Development and
Germination: Development of Dormancy; Germination. Weeds
and Competition: Gene Flow and Herbicide Resistance;
Herbicide Application Technology; Integrated Weed
Management.
Further Reading
Cousens, R., Mortimer, M., 1995. Dynamics of Weed Populations. Cambride University
Press, Cambridge.
Radosevich, S., Holt, J., Ghersa, C., 2007. Weed Ecology: Implications for Manage-
ment, third ed. John Wiley & Sons, New York.
Ross, M.A., Lembi, C.A., 2009. Applied Weed Science: Including the Ecology and
Management of Invasive Plants, third ed. Pearson Education, Inc., New Jersey.
Zimdahl, R.L., 2007. Fundamentals of Weed Science, third ed. Academic Press, San
Diego.