\HARAMAYA UNIVERSITY

POSTGRADUATE PROGRAMS DIRECTORATE

College: Agriculture and Environmental Science

School: Plant Science
Program: Agronomy
Course: weed science

Term paper:
Review on the study of Structure and Dynamics of weed population Using
Exponential and Logistic Growth Curve.

Prepared
by
Samuel Lemma
ID PGP/989/15

Submitted
to
Lemma Degebasa (PhD)

Haramaya University, Ethiopia

Jan 2023
Table of Contents

1.INTRODUCTION.................................................................................................................4

2. LITIRATURE REVIEW.....................................................................................................5

2.1 Over view of Weed Population Dynamics....................................................................5

2.2 Major Factors affecting Weed Population Dynamics.................................................5

2.2.1 Seed Pool Composition............................................................................................5

2.2.2 Cultural and Chemical Practices............................................................................6

2.2.3 Weed Population Changes due to Soil Property Modification............................7

2.3 The concept of Population Dynamics: Size Changes over Time................................8

2.4 Over View of Exponential and logistic growth curves................................................8

2.5 Real population growth curves....................................................................................11

2.6 Effects of Migration (Immigration and Emigration) on Population Size...............11

2.6.1 Migration among populations: creating Meta populations...............................12

2.7. Population Structure...................................................................................................13

2.7.1 Age structure..........................................................................................................13

2.7.2 Size structure..........................................................................................................14

2.7.3 Phenology................................................................................................................14

2.8 Why does population structure matter?.....................................................................15

3. Summary.............................................................................................................................16
 1. INTRODUCTION

A population is a group of individuals of the same species found in the same place at the
same time. Like many ecological terms, this definition is flexible, because it can be used to
describe populations at many scales. Similarly, a population may be the number of
individuals contained within a small area (e.g. a field) or it may refer to the local or regional
distribution of the species (Swanton, 2003). The first step in understanding any species is to
document its distribution and abundance. This gives the researcher an idea of the scope of the
potential problem (i.e. weediness). Note that we say potential problem: while distribution and
abundance are useful information, more data must be obtained before a decision is made on a
species’ weeding (González-Andujar, 2012).

Generally, populations were treated as whole entities. We then discussed the spatial
distribution of individuals within a population, and how this would influence estimates of
distribution and abundance. For the most part, we treated individuals as identical entities
(Navarrete, 2019). Populations, however, are made up of individuals that vary in age, size,
genetic structure (genotype) and appearance (phenotype). As a result, populations are
structured by this variation (Mortimer, 1995). Population structure refers to the organization
of individuals within a population, based on specific characteristics. For example, in a human
population we may compare the age structure of men and women.

Demography is the study of population size and structure, and how they change over time.
Populations are also dynamic: their size and structure change over time. Population size
refers to the total number of individuals or the density of individuals within a specific
population. A change in population structure will affect population dynamics; as population
size increases or decreases, the structure will be affected (Acker, et al 2020). Therefore this
term paper is conducted with the objectives to review population change over time using
exponential and logistic growth curve of weed population in general.
 2. LITIRATURE REVIEW

2.1 Over view of Weed Population Dynamics

The weed community within an agricultural area faces continual change from year to year as
species composition is modified and shifts. The dynamic population of a weed system is
influenced by natural factors, agricultural practices and their interactions ( Grime, J.P et .
1977) Understanding how these factors affect weed community structure, especially human
interaction, can better determine how future weed populations will be managed.

Weed population shifts are the changes in the individual organisms that make up the
population of a locality, often caused by changes in weed management practices. The bottom
line for a farmer, and an individual field, is that because weeds adapt they change. Those
practices that controlled weeds in your field this year probably won't provide acceptable weed
control in future years. Why? Because what you do today kills those weeds susceptible to
your management practices. The few weeds that survive today's management practices are
the parents of the weed problems you will have in the future: leaner, meaner, better able to
handle what you dish out.

This process has been going on for thousands of years. Every crop grown by humans, in
every field, in every year for the history of agriculture has resulted in the weeds you have in
your field today. Agronomic practices (e.g. mechanization, herbicides, new crops and crop
varieties) and an increase in the size and structure of land holdings can change the weed
species present (interspecific population shifts), as well as the genotypes of a species present
(intraspecific population shifts), in a local agroecosystem.

Inter-specific Population Shifts: Many weeds have undergone very large extensions of the
ranges (e.g. foxtails) they thrive in while others formerly widespread have all but
disappeared. Before the use of herbicides the weeds in Iowa corn fields were different than
they are today. At that earlier time broadleaf weeds were considered the major weed problem
(farmer’s grouped plants in their fields into "weeds" and "grasses"). With the introduction of
2, 4-D, selective weed control selected for the weeds that herbicides didn't kill. 2, 4-D killed
the broadleaf weeds in most cases, but did nothing to the grasses. All the foxtails and other
grasses suddenly became much larger weed problems than before. Giant foxtail was largely
unknown to mid-western growers before 1950. Since that time the same population shift story
has been repeated over and over again. In the last 10-20 years the appearance of wild proso
millet and woolly cupgrass, weeds largely unknown previously, have spread across the
countryside (Barrett, et al 1983).

Intra-specific Population Shifts: Selection for weedy adaptations can result in changes
within a species. The selection for new and better adapted variants of a species can cause
shifts in the composition of a population of an individual species in a field or area. Just as
herbicides selected and shifted different species in a fields weed community, herbicides and
other forces have caused shifts within populations of one species to better adapted variants
(Rees, et al. 1998)

2.2 Major Factors affecting Weed Population Dynamics

2.2.1 Seed Pool Composition

The most influential element in determining above-ground weed population make-up is the
composition of the soil seed bank, or the seed pool (Norris, et al.2007). The seed bank, or the
soil’s accumulation of weed seed within a specific site, is formed by seed rain within the
region from previous weed species that survived to produce seed as well as from seeds
dispersed into the region. It is from this reserve of seeds that the next generation of the weed
community will be produced. The seed bank remains the crux of weed population dynamics;
modifications to the existing seed pool, both natural and manmade, will directly affect
subsequent weed communities (Wallinga, et, al. (1995).

In natural systems, weed seed within the seed bank must overcome several obstacles in order
to remain viable for the next suitable growing season. Components of the seed pool face
mortality in a number of ways, including: consumption by organisms, aging of seed, or
germination and subsequent death in an unsuitable environment (Westerman et al., 2005).
Further persistence within the seed bank depends upon genetic components of the seed as
well as the conditions of the surrounding soil. Weed species that succeed at maintaining
sufficient quantities of seed within the seed bank will, most likely, be one of the dominant
species within the system and continue to contribute seed rain to the seed pool in future years
(Kelly, et al. 1996).

2.2.2 Cultural and Chemical Practices

In agricultural cropping systems, herbicides have historically been the predominant control
mechanism to keep weed populations in check. However, recent research has explored the
concept of manipulating the natural population dynamics of weed species with agronomic
practices and found these practices to be viable weed control tactics (Westerman et al., 2005).

Human influence over the varying numbers of weed species is limited by what determining
aspects that control weed populations can be modified. The factors that can be controlled or
altered by human interaction focus on increasing weed seed mortality and decreasing the
number of established weeds that can add further seed to the seed pool (Heesterbeek J.A.P.
2000). even though, Tillage of agricultural land targets the weed seed bank, and often reduces
weed seedling emergence due to seed burial or early germination in adverse conditions. In
more recent years, however, the adoption of conservation tillage has reduced the dependency
on tillage. In these reduced tillage systems, increased weed seed mortality may also be
achieved by allowing weed seed to remain exposed and vulnerable to greater predation (
Roland, J. et al 2007).

In either instance, the agricultural practice effectively modifies characteristics of weed

population dynamics in order to reduce the weed population within a region. Other
agronomic practices that have been shown to help control weed populations are crop rotation
and the timing of the planting date by rotating crops, farmers can control seed production
and/or plant establishment. (Story, et al. 1987) Continuation of a monoculture system selects
for weed species adapted to those particular management practices; rotation of crop choice
allows for a diversification of herbicide modes of action to be incorporated into a system and
disrupts the favourable environment of the predominant weed species in the system. Early
planting dates of crops not only offer the possibility of increased yield, but also allow the
crop valuable growing time without competition from weed seedlings (Gunsolus, et al.1994).
When weed seeds begin to emerge, crops have already become well established and are less
affected by seedling competition; weed seedlings remain at a disadvantage due to limited
resources and may not survive to produce seed. Late planting of crops also offers a benefit by
allowing weed seed to germinate and be destroyed prior to crop planting. Both rotation and
planting date achieve some manner of control over weed population size in an agricultural
area (Deevey, et al. 1947)

2.2.3 Weed Population Changes due to Soil Property Modification

Many opportunities exist to alter weed communities through agricultural practices by

modifying the properties of the surrounding soil. Presently, there are numerous ways
documented as to how soil characteristics can be altered to favor crop seed rather than weed
seed. Some alterations include: mulching, use of legume residues in lieu of synthetic
fertilizers, use of cover crops, and directed placement of soil amendments (Heesterbeek, et al.
2000).

Mulching, plastic covers, and cover crop incorporation can lend to suppression of weed
seedling emergence by reducing light penetration to the soil surface thus inhibiting seedling
growth. Mulches and plastic coverings are usually limited to gardens and high value systems
like plant propagation and tomatoes. For large agricultural productions, cover crops remain
the most widely used form of ground cover. The use of cover crops has also been reported to
reduce weed seedling growth through allelopathy, or chemical inhibition (Westerman et al.,
2005). Both practices make use of altering the growing conditions within or on the soil
surface in order to increase weed seedling mortality and reduce competitive pressure on the
crop.

Agricultural procedures that change the type and/or placement of nutrients and resources
have also been noted as effective control strategies against weed populations. The use of
legume-based green manures instead of synthetic fertilizers have shown the ability to provide
adequate nutrients for large seeded crops while the delayed nutrient release has been growth
inhibiting to small seeded weed species (Liebman, et al, 2000). Not only do these green
manures release nutrients essential for the crop, they also release potentially phytotoxic
chemical that can be harmful to germinating weed seed. If nutrients and resources like water
can be of limited availability to weed seed, then germinating weed seed will be reduced. This
limitation can be achieved by directing water and fertilizer placement into close proximity
with the crop and less available to surrounding weed seed (Corre, et al. (2013).

These alterations to soil amendments can modify soil properties to an extent that lessens the
impact of the weed community on agricultural settings. Weed populations are ever changing
and it is difficult to predict their future shifting patterns. Species composition within a region
is determined not only by the genetics of certain weed seeds, it is also dependent upon
external factors and stresses like weather conditions and presence or lack of seed predators.
When agricultural production is factored into the equation, there are many interactions that
can redefine the weed population composition within a region. It is these interactions and
their results that we must focus on better understanding in order to manipulate weed
population dynamics in our favour (Zhukova, L.A. (1980). Once we more fully understand
our role in the subtleties of weed population establishment, we can better employ multi-
faceted control measures that rely less and less on one dominant management system and
more on an integrated, sustainable weed management system (White, et al, 1970).

Fig 1. Population size changes over time

 2.3 The concept of Population Dynamics: Size Changes over Time

In nature, a population’s size will rarely remain constant. Within a short time frame,
population size may remain stable, steadily increase or decrease, or it may cycle regularly, or
in an unpredictable fashion. The rate of population change is dependent on the ratio of
individuals entering the population through births (B) or immigration (I) to individuals
leaving through deaths (D) or emigration (E). Thus, the change in a population’s size (N)
from one time period (t) to the next (t+1) can be represented by the equation: N(t+1) = Nt +
B – D + I – E.

Additionally Birth (or natality) is the addition of individuals to the population. For plants,
births may refer either to the number of seeds produced or seeds germinating or to individuals
produced via vegetative reproduction Mortality is the loss of individuals from the population
through death. Mortality rates and causes will change over time. In this review we look at
population growth curves, first using the exponential and logistic models of growth and then
by looking at real populations (Acker, et al. 2020).

2.4 Over View of Exponential and logistic growth curves

As long as births outnumber deaths (ignoring immigration and emigration), population

growth will be positive. Over generations, a population with a constant positive growth rate
will exhibit exponential growth. The greater the difference between birth rate and death rate,
the more rapid the increase (Roland, et al. 2007). The difference between birth rate and death
rate is the instantaneous rate of population increase (r). Therefore the exponential population
growth can be shown as:

dN/dt = rN or Nt+1 = Nt ert

Where dN/dt is the change in N during time (t).

Many plants have the potential to produce a huge number of offspring. This is especially true
for some weeds where a single individual may produce more than 1,000,000 seeds per season
Given that plants produce so many seeds, why then do their populations not continue to
increase exponentially? Many seeds will not be viable, while others will not germinate
because environmental conditions are not appropriate, or because the seed dies due to
predation or disease. In spite of this, there can still be many viable seedlings produced per
adult plant. During the early stages of population growth, density may increase exponentially
but at some point, the growth will slow and density may even begin to decrease. Why is this
so?
Exponential growth cannot be maintained because populations are limited by a lack of
resources. At some point there will not be enough resources (e.g. nutrients, light or space) to
satisfy the needs of every new individual and so population density will level off.

The logistic curve is a model of population growth under limiting resources. Once a seed
germinates, there are many biotic and abiotic forces that cause mortality and reduce
population growth rate. For example, each seedling requires resources (space, nutrients,
water, light) to survive, and individuals that fail to acquire adequate resources will fail to
reproduce or may die. The lack of adequate resources will cause the population growth curve
to level off. The growth of all populations will eventually level off. The carrying capacity (K)
is the maximum number of individuals the environment can support. To incorporate K into
the population growth equation, the exponential equation can be modified by including an
additional term that causes the growth rate to level off. It looks like this:

dN / dt = r N dN/dt = rN (K-N)/k

Fig 2 exponential growth curve. Fig3 logistic growth curve.

logistic growth-curve equation which incorporates limits to population growth over time.
When population density (N) is less than K, the term (K-N)/N will be positive and population
growth will be positive. As the value of N approaches K, the rate of growth decreases until
N=K when the rate of population growth (dN/dt) becomes zero. The population size is stable
because births equals deaths at this time (Hulme, et al. (2005).

There are three parts to the logistic growth curve. Initially, population size increases at an
exponential rate. The maximum rate of growth occurs at half the value of K. Beyond this, the
rate of population increase slows down but is still positive (Ullman, et al. 1998).
This occurs because not all individuals will be affected by limiting resources at the same time
because of differences in size, age, health and reproductive status. Over time, the proportion
of individuals affected by limiting resources will increase and this causes the curve to level
off at K.
2.5 Real population growth curves

The exponential and logistic growth curves are idealized mathematical descriptions of how
population size will change over time. They provide a conceptual framework on which to
base more complex approaches to population growth. In real situations, population growth is
more variable over time. There are a number of reasons why population size fluctuates over
time.
 In the logistic growth model assumes that the environment is stable over time and
therefore K remains stable. This is unrealistic because the abiotic environment is
naturally variable: temperature, nutrients, water and light change over time. Even
small changes in one factor can affect the number of individuals the environment can
support.
 there is random variation in birth and death rates. This is termed demographic
stochasticity. An occasional low birth rate or high death rate can cause the population
to become extinct. Additionally The logistic and exponential growth curves assume
that populations are independent of other populations. Populations, however, interact
(through competition, herbivore) and this causes population size to fluctuate.
 The logistic and exponential growth curves assume that populations are independent
of other populations. Populations, however, interact (through competition, herbivore)
and this causes population size to fluctuate

2.6 Effects of Migration (Immigration and Emigration) on Population Size

Sometimes it may be possible to ignore the effects of immigration and emigration (migration)
by assuming that they are equal, or that their effect on population size is negligible. However,
many will be dependent on the immigration of individuals from other populations. A
population with fewer births than deaths will remain viable only when supported by seeds
imported from other populations (Grace, et al. 1991).
Migration demographically links populations. Determining whether migration is an important
demographic process has two problems. First, the concept of migration assumes that there are
specific boundaries over which individuals (seeds) move. In human populations we have
political boundaries, so we can keep track of the movement of (most) individuals. Plant
population boundaries are rarely discrete. Second, even if ‘real’ boundaries do exist, tracking
the movement of individuals can be challenging. Therefore, it is difficult to establish if
migration is occurring.

2.6.1 Migration among populations: creating Meta populations

Traditionally, populations have been described as a collection of individuals that are capable
of interbreeding. In reality, most populations are scattered and clustered into smaller
subgroups. This clustering may be a random process but it usually reflects the heterogeneity
of the landscape, i.e. there are a limited number of areas where individuals of various species
can live and these individuals cluster in amenable habitats. When populations become
divided into clusters, we can say that each cluster becomes spatially isolated from each other.
If spatially isolated populations interact through migration (e.g. of seed) or distant
pollination, then the aggregate of interacting populations is called a ‘meta population’. The
implication of using the term ‘meta population’ is that interactions among populations are not
always common but they do occur.

Each population within a Meta population will likely be genetically distinct because each is
adapted to local environmental conditions. Although individuals within a population will
mostly mate with individuals from their own population, Meta population dynamics will
introduce some genetic material from surrounding populations. Since the continued existence
of a population is determined mainly by whether there is enough local genetic variation to
withstand environmental change (including diseases, herbivore, drought) and ensure births
exceed deaths, Meta population dynamics may prevent the extinction of local populations.
For example, immigrants (or at least their genetic material via pollen) from other populations
can help maintain a population that otherwise would become extirpated (locally extinct)
because it is not genetically suited to changes in its environment (e.g. decreasing light levels).
Populations that are maintained only through immigration from other (source) populations are
called ‘sink’ populations (Pulliam, et al, 1988). In weedy white Campion (Silene alba), for
example, isolated populations survive only because new genetic material arrives via
immigration from surrounding populations – in this case, the immigrant genetic material is
delivered via pollen (Richards, et al, 2000)

Perhaps the most important aspect of the meta population concept is the implication for
conservation. Because populations may contain relatively few individuals, be
Restricted to a small area or have low genetic variation, they are subject to local extinction.
However, the Meta population is usually persistent because local adaptations In populations
increase the total amount of genetic variation. If the landscape-scale environment changes
suddenly, the chances are good that at least one population has the genes needed to allow for
decolonization of habitats vacated by local extinctions. This means that should a disease or a
drought strike, then some of the populations will survive. Over time, this means that the local
habitats that populations occupy often are ‘emptied’ and recolonized many times.

Therefore while local populations may go extinct and the habitats emptied, the Meta
population of a species will continue. This has become important in understanding how to
conserve species. It is possible that a large contiguous reserve that does not allow for spatial
isolation, local adaptation, and development of a Meta population can actually hasten
extinction of a species, as it is vulnerable to sudden environmental change (Beeby, 1994;
Hanski and Gilpin, 1997; Schwartz, 1997; Honnay et al., 1999; Etienne and Heesterbeek, et
al, 2000).

2.7. Population Structure

Populations are characterized based on the age, size, appearance or genetic structure of
individuals. In fact, population structure could be based on any characteristic that is variable
within a population. Population structure is not a static feature of a population because
individuals age, grow, reproduce and die at different rates depending on their individual
characteristics and their environment. In this chapter, we focus on age, size and
developmental stage structure of populations.

2.7.1 Age structure

The distribution of ages within a population can be characteristic of the species itself, or it
can reflect the ‘health’ of the population, or the environment inhabited by the population. In a
‘healthy’ population, younger individuals will outnumber older individuals because a
proportion of young individuals will die before they reach maturity. (Whipple, et al (1979)
proposed five age class distributions to explain population trends of trees. The ‘inverse-J’
curve shows a population with many more juveniles than adults; this population is likely to
be relatively constant or increasing. The ‘bimodal’ distribution is a result of pulse recruitment
(addition of new individuals) where periods of lower recruitment are followed by periods of
higher recruitment. This population will likely be stable or increase as long as recruitment
pulses are frequent enough to replace dying individuals. A ‘decreasing’ population
distribution means the population is not replacing itself because recruitment is not high
enough to replace those that are dying. If recruitment is zero the distribution will become
‘unimodal’ as the population ages and no young individuals are added. Although individuals
are present, the population will become extinct unless increased reproduction occurs. Finally,
a random distribution is typical of a population in a marginal habitat, or one that is
responding to disturbance. Populations that have recently invaded a site are also likely to
exhibit this distribution (Luken, et al, 1990).

Age structures can be difficult to interpret because they do not always fit the theoretical
distributions described above, nor are they consistent over time. Montana populations of
spotted knapweed (Centaurea maculosa) tended to have inverse-J distributions in 1984, but in
1985 the distribution decreased. This occurred following a severe drought in 1984, when
young individuals experienced higher mortality than older individuals (Boggs, et al, 1987).

2.7.2 Size structure

Most populations will tend to have fewer large individuals and many smaller ones. However,
larger individuals can have a disproportionate effect on the rest of the population because
they tend to live longer and produce more offspring than smaller individuals of the same age
(Leverich and Levin, 1979). Larger individuals can also directly affect smaller individuals
through shading. Plant size is a measure of the success of an individual because the larger
individuals have acquired more resources than smaller individuals. For this reason, it is often
more useful to structure populations by size rather than age. Furthermore, size may be a
better predictor of an event (e.g. reproduction or death) than age (Werner, 1975; Werner and
Caswell, 1977; Gross, 1981).

2.7.3 Phenology

A plant’s phenology (stage of development) can be used in conjunction with or instead of

plant age and size to examine population structure (Sharitz and McCormick, 1973; Werner
and Caswell, 1977; Gatsuk et al., 1980; Horvitz and Schemske, 1995; Deen et al., 2001). This
measure may be more biologically meaningful than age or size alone because an individual’s
Phenological stage may be more linked to its likelihood of survival or reproduction. Horvitz
and Schemske (1995) showed that the annual survival and fertility of the prayer plant
(Calathea ovandensis), varied depending on the individual’s Phenological stage. Seedlings
had less than 10% survival, seeds and juveniles had moderate survival while other stage
classes had over 90% survival. Reproductive individuals produced different numbers of seeds
per plant depending on their size.

2.7 Why does population structure matter?

Interpreting population structure can be difficult and time consuming. Why, then, do we do
it? Why not simply calculate population means (e.g. mean age or height) and use these simple
numbers to describe a population? The answer is that there is a lot of valuable information in
the variability of a population and that by reducing this to a mean value we lose information
(Hutchings, 1997). Populations may have the same mean values but different structure have
the same mean stem diameter(x=20 cm) but the proportion of individuals in each size class
differs. The fate of these populations will most likely differ. Another reason we may be
interested in the structure of a population is that it is away to identify specific individuals of
interest. For example, we may only be only interested in plants of a certain size (age or
stage). If we know, for example, that only individuals above a specific size will impact crop
yield, then we can focus our research on the larger age classes. Recognizing population
structure helps us to focus on specific individuals within a population, and it gives us a
glimpse of possible population dynamics to come.

3. Summary

Describing population dynamics, population structures is difficult because of genetic and

environmental variation and the complex interactions and combinations that can occur. This
complexity is the reason why our convenient measures and descriptions of populations are
often not adequate even if they do a reasonable job of approximating the real world. This
complexity explains why:

 simple logistic and exponential equations do not adequately describe populations;

 spatial isolation within meta populations influences survival and conservation
decisions;
 classifying plant population structure by age, growth stage, size and life cycle can be
difficult;
 life history strategies are good rules of thumb but not all that accurate in predicting
the population dynamics and impact of plants, especially weeds.

Population dynamics and structure are good concepts to understand, but they need to be
developed and studied in the context of ecological interactions and genetic variation. This
means it is not enough to understand the general patterns of populations. We should also
understand how populations change with genetic diversity, variation in reproduction, and
with the presence of competitors, herbivores and disease.
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