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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 What Traits Make Plants a Weed?

What Are Harmful and Beneficial Aspects of Weeds?
In this article, weeds are defined and the common features of
their biology described. Population dynamics of weeds is Inherent in the definition of a weed is the idea that it is harm-
considered and a brief outline of options for managing weeds ful, to other crops, humans, livestock, and the environment.
is given. 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
What Is a Weed? weeds, they would not be classified as such and could provide
benefits. Losses due to weeds can be categorized as direct and
A simple definition of a weed is a plant growing where it is not indirect losses. Direct losses would include those that reduce
wanted. As humans, we have decided which plants are desir- both the quantity and quality of the ‘crop’ being produced.
able in a given location, and which plants are detrimental to These losses are associated with weed presence in any agricul-
our desired goal. In a crop production situation, the presence tural production situation, such as row crops, vegetable and
of weeds can have a harmful effect on the desired crop’s yield fruit crops, ornamental and trees, turf and athletic fields,
or quality. In a flower bed or vegetable production situation, pasture and rangeland, or natural environments. Indirect losses
presence of weeds can detract from the esthetic value, make it would include those that do not reduce cash return from the
difficult to harvest the vegetables, and reduce the quality of crop being produced, but represent a cost to society at large
the produce. The Weed Science Society of America defined or to the producer or property owner. For example, harboring
a weed as “a plant growing where it is not desired.” The Euro- insects, diseases and pests of other crops, creation of health
pean Weed Research Society uses the following definition: “a hazards such as pollen from common ragweed that results in
weed is any plant or vegetation, excluding fungi, interfering hay fever suffering in humans or dermatitis by poison ivy
with the objectives or requirements of people.” leaf toxins; reduced property values due to weed invasion,
A weed is often a plant out of place, such that a corn plant and cost of weed maintenance in noncrop environments,
growing in the field where it was planted is desirable, but when rights-of-way, public lands, and so on.
it appears in a soybean crop the following year, it is considered The main measure of harm is in lost yields. Increasing weed
a weed. Thus, any plant species and any type of plant can be densities in row crops cause increasingly greater losses of
considered a weed. It is very difficult to describe all traits of yields. Even with adequate weed control efforts, weed presence
all plants that might be considered weedy. can still cause significant loss (10–25%) of grain yields. The
Weedy plants can be found in very different environments next main harmful measure is reduced quality of harvested
across different taxonomic classifications, including aquatic, product that could be due to contamination by weed
terrestrial, and aerial environments. In aquatic environments, seed (jointed goatgrass in winter wheat), weedy plant parts
blue-green algae and macrophytes such as elodea (Elodea cana- (Canada thistle heads in canned garden peas), seed staining
densis) can be troublesome plants. In aerial environments, (eastern black nightshade in soybean). Often this contamina-
bromeliads such as airplants (Tillandsia) take advantage of tion results in extra costs for cleaning the product or dockage
stressed trees and open leaf canopies to grow on tree branches. for the crop when sold. Presence of weeds can result in harvest-
Most weedy plants occur in terrestrial environments such as ing difficulties, such as remaining green while the crop senes-
summer annual broadleaf and grass weeds in crop fields, peren- ces prior to harvest and not able to be cut and processed
nial and brushy weeds in range and pasture lands, dandelion or through a combine harvester or other such machine. If weeds
mosses in urban lawns or sports turf, mosses or liverworts in are present during hand-harvesting of vegetable and fruit
ornamental nursery pots and creeping perennials such as field crops, it interferes with efficiency. Additional costs include
bindweed or quackgrass in gardens. Taxonomically, plants processing of harvested crop and/or cost of transporting
considered weeds could be algae, mosses and liverworts, ferns weed seed with grain, or loss of storage efficiency with green
and horsetails, to gymnosperms (eastern red cedar or salt plant parts in harvested crop.
cedar), or angiosperms (flowering plants). Weeds are known to harbor insect and disease pests as alter-
What is the biology of weeds? By knowing unique charac- native hosts and as a ‘green bridge’ between different crops in
teristics for an individual plant species, we can understand a rotation sequence to increase pest management costs,
where will it grow well based on a location’s environmental possible yield losses, in a subsequent crop. For example,
conditions of temperature, moisture, and day length; we can numerous soybean pests use weeds as alternate hosts, such as
use information about its lifecycle (i.e., annual, biennial, or Asian soybean rust on kudzu, soybean aphids on buckthorn,
perennial) to know when it will occur in a growing season and soybean cyst nematodes on winter annual weeds such as
and how long it will survive; does it reproduce by seed or purple deadnettle and henbit. Presence of green foxtail
vegetatively or both, and how does it spread from location provides a bridge between winter wheat crops by harboring
to location. aphid pests that transfer viruses.

Encyclopedia of Applied Plant Sciences, 2nd edition, Volume 3 http://dx.doi.org/10.1016/B978-0-12-394807-6.00026-5 469

470 Weeds and Competition j Weed Biology

There are a few weedy plants that are parasitic on beneficial pennycress (Thlaspi arvense) for oil or switchgrass (Panicum vir-
plants and can result in reduced crop growth and yield reduc- gatum) for cellulosic biofuel, used for coloring and dyes, and
tions, such as field dodder (Cuscuta spp.) in alfalfa or clover a component of biodiversity in agricultural areas. Many crop
crops; broomrape (Orobanche spp.) which attacks carrots, fields in North American agriculture are monocultures of row
tomatoes, sunflowers, redclover, and witchweed (Striga spp. crops such as maize and soybean, thus described as having
mainly S. hermonthica) that can infect corn and sorghum and reduced biodiversity. In other places, policies are in place to
drastically reduce their grain yields by 10% compared to encourage farmland diversity either by reducing intensity of
a healthy crop. Dwarf mistletoe (Arceuthobium spp.) is a parasite management or by reducing the farmed area through set aside,
on pines and firs while broadleaf mistletoe ( Phoradendron mac- such as a current European Union policy. By these practices,
rophyllum) infects numerous landscape trees; each reduces weedy plants contribute to increased biodiversity.
wood quality, increases chance for disease infection, and results There is interest in designing weed management systems that
in death of the trees. could enhance the beneficial aspects of weedy plants, for
Presence of weeds in aquatic environments can prevent example, that would allow for development of populations of
water flow by blocking ditches and irrigation canals, and tran- beneficial organisms, such as pollinating insects, seed-eating
spire water that could have been delivered to crop plants. The birds, insects, and mammals, and harbor natural predators,
water hyacinth ( Eichhornia crassipes) is an important tropical such as ground beetles and spiders, of aphid pests of crops.
aquatic weed. Eurasian watermilfoil ( Myriophyllum spicatum) For example, if a certain level of weeds in a crop could be toler-
is considered to be originally from Europe and Asia, released ated, this may allow for more farmland birds to survive and to
into North American lakes and rivers and has drastically altered rear chicks before these food sources are removed. Weedy plants
the ecology of infested water bodies. The sheer mass of plants in stubble fields provide a level of erosion control from wind
can cause flooding, and the stagnant mats are good habitat and water if there is little residue. However, successful weed
for mosquitoes. Milfoil mats can rob oxygen from the water control practices would still need to be available prior to estab-
by preventing the wind from mixing the oxygenated surface lishing a crop in these fields. Concerns would be raised if there
waters to deeper water. The dense mats of vegetation can also was significant presence of herbicide-resistant weed populations
increase sedimentation rates by trapping sediments. Milfoil allowed to grow uninhibited.
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 What Are the Population Dynamics of Weeds?
weeds may reduce oxygen content of the water and lead to
fish deaths. Weed species can be categorized into different life cycles,
Harmful aspects of weeds on humans include discomfort including annual (winter or summer), biennial, and perennial
due to allergies causing ‘hay fever’ (pollen from trees, grasses, (simple, creeping, or woody). Important features of each life
and weeds such as common ragweed (Ambrosia artemisiifolia) cycle include time of seed germination, number of seedlings
and due to dermatitis caused by the oily sap from poison ivy established, rate of vegetative growth, time of flowering and
(Toxicodendron radicans), poison oak (T. diversilobum), or seed production, and seed dispersal. Annual weeds complete
poison sumac (T. vernix) that contains urushiol and when on their life cycle within a period of 12 months or less, while bien-
skin, results in an itchy rash anywhere from a few hours to nial weed species require up to 24 months, and perennials
days after exposure. survive longer than 2 years, sometimes not producing flowers
There are many weed species that impact animal production and/or seed for several years until plants are well established.
and animal health. Animals can die from eating poisonous Perennials often have the capability of regenerating from vege-
plants, off-flavor is imparted to milk and other dairy products tative structures (roots, adventitious buds, tubers, rhizomes,
when weeds such as wild garlic (Allium vineale) are eaten, and stolons, and so on) in addition to seed.
livestock hides and carcasses are physically damaged by weeds.
Plants with spines or thorns, such as thistles, inhibit animal
Weed Seed Dispersal in Space and Time
foraging and protect neighboring plants from being eaten,
thus reducing the area available for grazing by livestock. The biology of annual and biennial weed species emphasizes
Some animals, such as horses, are much more sensitive to reproduction by seed. Very high levels of seed production per
certain plants than others, and if these plants are present in plant are common along with unique dispersal mechanisms
degraded pasture areas so that they are grazed by horses, such in space via wind, water, animals, and humans, as well as
as poison hemlock (Conium maculatum), tansy ragwort (Senecio dispersal mechanisms in time via dormancy. Seed dispersal
jacobaea), yellow starthistle and Russian knapweed (Centauria by plants in space is a passive process, in the sense that plants
spp.), they can experience chronic effects if enough is cannot actively choose where their offspring will go. However,
consumed, and result in animal death. various features of the seed itself and the maternal plant can aid
It may seem strange to consider the positive effects of weeds; dispersal. Taller plants may disperse seeds over greater
however, weeds are just plants and only from human perspec- distances. Lighter seeds may be dispersed further by wind,
tive, are they troublesome. Beneficial aspects of weeds include but may have a lower chance of becoming a new seedling
pollination and nectar resources for bees, monarch butterflies, because of limited stored energy reserves. Heavier seeds fall
and other insects, source of genetic traits that could be intro- near the maternal plant, where they may have a greater chance
gressed into crop plants for improved fitness, biomass produc- of landing in a favorable site for germination since it was
tion that could be used as biofuel resources such as field a successful location for the parent, but could be detrimental
Weeds and Competition j Weed Biology 471

due to future competition with siblings or increased likelihood the reinvasion rate. These factors can lead to large-scale
of predation by seed predators. In general, most seed fall near temporal and spatial variations in overall and local population
the parent plant if no active mechanism is assisting the sizes.
dispersal. To achieve greater distances, some seed or fruit con-
taining the seed can be moved internally by animals or because
Weed Introduction (to New Sites)
of unique appendages moved externally when attached to the
animal. For the weed species, the advantage of dispersal in Seed dispersal by plants is a passive process, but various plant
space is to find new sites, while a disadvantage might be the and seed features aid in their dispersal. The goal of seed
absence of another plant needed to achieve cross pollination dispersal in space is to seek out new and better environments
for successful seed production. The establishment of a soil for future generations, to aid in genetic mixing among indi-
seedbank provides a means for the weed species to have viduals, and to not be competing with parent plant or future
different generations and genetics that can appear in future ‘siblings.’ A drawback may be the absence of another related
years (see article Weeds and Competition: Weed Seed Biology plant with which to achieve cross pollination for production
for more details on seed rain, soil seedbank, and seed of new seed. Individual weed species produce differing
dormancy/germination). 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,
Modeling the Population Dynamics of Weed Species
plants could produce many lightweight seed to be dispersed
Understanding the overall dynamics of a weed population widely by wind as compared to few heavyweight seed that
provides insights for making management decision based fall near the parent plant. Taller plants may disperse seed
on when the population might be most vulnerable, but not over greater distances from the parent plant. Many seeds
on an expectation that the measure will be successful. Often, also have adaptations for animal dispersal (internal or
the seedling growth stages are targeted for weed control early external to the animal) that encourage even greater dispersal
in the cropping season, but if these seedlings only represent distances. Fleshy fruits are aided by birds or other species
a small fraction of the total soil seedbank, another aspect of that eat the fruit and transport the seed long distances before
the population should also be targeted, such as the amount being deposited. Externally, burs and seed with sticky coats
of seed produced by mature plants and subsequently returned attach to animal fur or human clothing and are transported
to the seedbank each year, or by manipulating the placement until rubbed off.
of seed in the seedbank to reduce their longevity. Many of the (For features of ‘seed rain,’ the seedbank in the soil, and the
models of weed populations are based on information on topics of seed dormancy and germination, see Weeds and
reproduction of a limited number of weed species and are Competition: Weed Seed Biology.)
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 Managing Weeds
of management decisions (concerning use of tillage practices,
time of crop planting, selection of crop type, management of Weed management is covered in more detail in the article. A
fallow periods, and so on) in alternative scenarios, but inevi- successful weed management strategy does exploit our knowl-
tably can only be as good as the data on which the models are edge of weed biology to achieve crops with minimal presence
based. Models can be used to assess the efficacy of interven- of weeds, levels known to not affect crop productivity beyond
tion at different stages in the life cycle of weeds. They may the economic cost.
assist in decision making on whether to use a cheaper herbi- Weeds, like crops, are living organisms that respond to their
cide that is less effective, rather than on a more effective but environment. We manipulate the environment to produce
more expensive one. Models can help in deciding whether conditions conducive for growth of crop plants, but weeds
or not to treat the weed, that is to incorporate a weed also exploit these optimal conditions. The optimal plant envi-
‘threshold’ into the management decision in order to prevent ronment should give the advantage to the crops over the weeds.
unnecessary or uneconomic treatments. Weed management is one of the key considerations among
If weeds have been managed well in an intensive agricul- many for successful crop production, and each decision and
tural system, individual species often show patchy or scattered choice made influences the likely weed species and infestation
spatial distributions among fields. Weed management by some levels that will be observed in their fields. The time of the
farmers might lead to local extinction of a given weed species previous harvest, management of the residues of the previous
(but often difficult because of the soil seedbank and persistent crop, type and timing of soil tillage, fertility management,
seed for some species). However, reinvasion can easily occur choice of crop species and variety, date of planting, and row
due to dispersal of seed by wind, water, animals, and/or spacing and seeding rates are all decisions made about crop
humans. The small, sometimes transient, populations in a land- production that will influence the type of weed population
scape might be termed a metapopulation. The dynamics of the that appears in the field.
metapopulation (multiple small patches that are intercon- An integrated approach to developing weed management
nected via dispersal) will depend partly on the size of the local systems is recommended, and this includes knowledge of the
populations or patches, and their rate of extinction. The biology of the weed species occurring in the field, determining
connectivity among patches is important and determined by the level of infestation (is management even necessary?), and
dispersal rate and distance between patches that can determine then evaluating numerous options. These control tactics can
472 Weeds and Competition j Weed Biology

be categorized as cultural, physical, biological, and chemical repeated use of single herbicides or single herbicide modes
options. of action has led to the evolution of herbicide-resistant weed
Assessing weed populations in the field can be problematic. biotypes, which have proved challenging to manage.
First step is correct identification of the weeds species, and this Continued focus on understanding weed biology and use of
should be done at the seedling stage of growth if timely control integrated weed management systems should provide effective
has to be implemented. The distribution of weeds in a field means of managing these plants.
tends to be patchy, thus requiring the entire field to be scouted.
The implication of weed patchiness is that the application of See also: Arable Crops: Agricultural Crops; Agroecology:
control can be localized rather than across the entire field. If Farming Systems with Nature as Guide. Plants and the
infestation levels are relatively low, it will be inappropriate to Environment: Invasive Plant Species. Seed Development and
determine an average infestation for a whole field that results Germination: Development of Dormancy; Germination. Weeds
in underestimating weed populations in high-density patches and Competition: Gene Flow and Herbicide Resistance;
and recommending weed control in weed-free or low-density Herbicide Application Technology; Integrated Weed
areas. Management.

Weeds in the Future

Further Reading
Weed species are very adaptable. Changes in crop management
and in climate are likely to alter the occurrence of weed species
Cousens, R., Mortimer, M., 1995. Dynamics of Weed Populations. Cambride University
in fields. Climate change may result in the shift of weed species Press, Cambridge.
occurrence so that ‘new’ weeds from other areas move either Radosevich, S., Holt, J., Ghersa, C., 2007. Weed Ecology: Implications for Manage-
as a direct response or as a response to changes in cropping ment, third ed. John Wiley & Sons, New York.
systems. Invasions of new species into areas will generate 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.
new challenges for growers. Our global economy often encour-
Zimdahl, R.L., 2007. Fundamentals of Weed Science, third ed. Academic Press, San
ages movement of plant species around the world, and thus Diego.
potentially introductions to new areas. Reliance on the