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Sustainable Agriculture Reviews
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ISSN 2210-4410 ISSN 2210-4429 (electronic)

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DOI 10.1007/978-94-007-5961-9
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Contents
Surprising Facts About Soils, Students and Teachers!

A Survey of Educational Research and Resources ...................................... 1
Douglas Hayhoe
Sustainable Agriculture and Climate Changes in Egypt............................. 41
Hassan R. El-Ramady, Samia M. El-Marsafawy, and Lowell N. Lewis
Adapting Maize Crop to Diverse Agro-Ecosystems ..................................... 97
Ioannis S. Tokatlidis
Intercropping ................................................................................................... 119
Ehsan Neamatollahi, Mohammad Reza Jahansuz, Dariush Mazaheri,
and Mohammad Bannayan
Enhancing Fertilizer Efficiency in High Input
Cropping Systems in Florida ......................................................................... 143
Johannes M.S. Scholberg, Lincoln Zotarelli, Michael D. Dukes,
Monica Ozores-Hampton, Guodong Liu, and Pablo Tittonell
Soil Organic Matter Dynamics and Structure ............................................. 175
Nikolaos P. Nikolaidis and G. Bidoglio
Plant and Animal Breeding as Starting Points
for Sustainable Agriculture ............................................................................ 201
Gerhard Flachowsky, Ulrich Meyer, and Manfred Gruen
Medicinal Plant Active Compounds Produced
by UV-B Exposure ........................................................................................... 225
Rima Kumari and Majeti Narasimha Vara Prasad
v
vi Contents
Assessing the Environmental Benefits of Compost

Use-on-Land through an LCA Perspective................................................... 255
Julia Martínez-Blanco, Cristina Lazcano, Alessio Boldrin,
Pere Muñoz, Joan Rieradevall, Jacob Møller, Assumpció Antón,
and Thomas H. Christensen
Biocontrol of Plant Pathogens Using Plant
Growth Promoting Bacteria ........................................................................... 319
Pratibha Prashar, Neera Kapoor, and Sarita Sachdeva
Index ................................................................................................................. 361

Surprising Facts About Soils, Students
and Teachers! A Survey of Educational
Research and Resources
Douglas Hayhoe
Abstract Soil is one of the key resources that sustain life on Earth, not just as the
foundation for almost all our food supplies, as important as that is, but also in the
way that it filters water, supports biodiversity, and perhaps even moderates global
climate. Yet the world’s soils are under increased pressure on many fronts. They
face unprecedented threats from erosion, deforestation, desertification, salinization,
sealing (paving over), contamination, loss of biodiversity, and climate change. The
importance of soil and the need to sustain it against these threats, however, have
elicited little interest, not only by scientists and the general public, but also by
the educational systems of most countries. While increasing attention has been paid
to other important environmental topics, such as loss of biodiversity, climate change,
deforestation, fresh water availability, and the world’s oceans, little attention has
been placed on soil so far.
A way of meeting this challenge that has been instituted in a few countries has
been to include soil science, e.g. its concepts, concerns and protection, as a core
topic in the country’s national science curriculum, so that from a young age students
learn the key concepts of soil science and how and why people should protect soil
in a sustainable way. The research surveyed in this article shows that elementary
students as young as preschool have some initial ideas about the depth of soil and
its usefulness in supporting plant growth, but have little understanding of its com-
position, formation, or origin. Middle school students, of 10–12 years in age, arrive
at the topic with more understanding in some areas, such as the thinness of soil layers,
but are still ignorant concerning its age and origin. After several weeks of hands-on
activities combined with “minds-on” discussion, students as young as 5–6 years in
age are able to get “soil on their mind,” as evidenced by the diagrams they draw before
and after intervention, while students 10–12 years in age are able to understand
D. Hayhoe (*)
Department of Education, Tyndale University College,
25 Ballyconnor Court, Toronto, Ontario M2M 4B3, Canada
e-mail: dhayhoe@tyndale.ca
E. Lichtfouse (ed.), Sustainable Agriculture Reviews, Sustainable Agriculture Reviews 12, 1

DOI 10.1007/978-94-007-5961-9_1, © Springer Science+Business Media Dordrecht 2013
2 D. Hayhoe
the three-dimensional nature of soil, as well as start to understand its formation

process and age.
Elementary teachers begin their profession understanding substantially more soil
concepts than their students. Over 80% know that soil is formed by the weathering
of rocks, that earthworms mix the soil and allow for more air and water to enter, and
that decomposition provides soil nutrients for plants to grow. Very few of them,
however, are aware of how many life forms there are in a handful of soil, how many
years it takes for soil to form, how much of soil is space for air and water, which
component of soil has the smallest particles, or what role humus plays. After two or
three classes of intensive hands-on activities, they also are able to make substantial
gains in their understanding, reducing by 33% what was lacking in their understand-
ing of soil concepts. They can also make gains in their attitudes towards the need to
protect soils, compared with other environmental challenges.
The little research that has been done with secondary students shows that their
initial ideas about soils, and their ability to achieve a deeper understanding of soil
through classroom activities, is similar to that of middle school students. No studies
have reported on secondary school science teachers’ understanding of soil. Two
studies with secondary school agriculture teachers indicate mixed results as to how
prepared they are to teach soil science. This review concludes with a brief descrip-
tion of resources available for soils education, including equipment kits and unit
manuals for elementary school, and journal articles, websites, and electronic
resources for all grades. Given available soil education research and resources, this
work suggests that the most important thing people concerned about soil education
can do is advocate for the inclusion of soil science as a separate topic in their national
elementary science curriculum, if that is not already in place.
Keywords Soil concepts • Sustainability • Education • Hands-on activities • Initial

ideas • Gains in understanding
1 Introduction
Soils are one of the planet’s most important and indispensible resources.
Understanding soils is key to properly sustaining them. Given the need to feed a
growing world, there is a great deal of research that focuses on the role of soils in
agriculture (Banwart 2011). Healthy soils, however, are not only essential for food
and forests; they also filter water, transform nutrients, and sustain the world’s biodi-
versity. Furthermore, according to soil researcher John Zak, they may also play an
important feedback role in climate change projections (personal communication,
April 24, 2012). Yet of the world’s most basic resources, soils remain the least
studied and the least understood, both among scientists as well as the general public,
although a recent focus on soil awareness and education by soil scientists in Europe
indicates an ability by people of all ages, starting with young children, to learn key
concepts about soil science (Fig. 1).
Surprising Facts About Soils, Students and Teachers… 3
Fig. 1 Young children investigate a soil profile in Greven, Germany, during “Soil Action Week”
(County Steinfurt 2010). The European network on soil awareness helps organize soil awareness
public events at various locations in different countries (Broll 2011)
Science education research – students’ initial understandings of science concepts,

the most effective teaching strategies, etc. – has been blossoming for three decades,
with over a thousand conference papers and peer-review journal articles appearing
annually. Very little research on elementary and secondary soil science education,
however, has been reported. For example, the initial understandings and misconcep-
tions of school children or their teachers about light and what pedagogical strategies
are most effective in helping students learn about light have been the focus of at
least 50 published articles. A similar abundance of information is available for many
other school science topics such as force, motion, electricity, matter, substances,
chemical reactions, plants and animals, ecosystems, the cell, and reproduction. Even
in the Earth and Space section of the science curriculum, topics much less crucial to
our survival than soils science– earthquakes and volcanoes, rocks and minerals,
phases of the moon, stars and planets – are mentioned much more frequently.
In contrast, perhaps one or two articles are published on elementary and secondary
soil education each year.
Soil science does not feature prominently in most educational systems, at least in
America. At the University of Florida, for example, a campus with almost 50,000
students, the College of Agriculture and Life Sciences has some 4,500 students; but
only ten of them are enrolled in soil and water science (Collins 2008). This study by
4 D. Hayhoe
Table 1 The role of soil in elementary and secondary science curriculum documents in some
English speaking countries and regions compared with other earth and space topics
Australia (ACARA 2012) Soil is briefly referred to as one of earth’s resources in
Gr. 2 and as part of the changes in Earth’s surface
over time in Gr. 4. Other earth and space topics,
however, are given more prominence. Topics in
astronomy, for example (sun and moon, solar
system, and stars), are addressed in Gr. 1, 3, 5, 7, and
10; and topics in geology (rocks and minerals, plate
tectonics, and natural disasters) are addressed in
Gr. 4, 6, 8, and 9
California (CDE 2009) Soil is mentioned several times in Gr. 2 Earth Sciences,
with rocks, the rock cycle, and erosion, and several
times in Gr. 6, with topography, ecosystems, and
natural resources. Topics in astronomy and geology,
however, such as rocks, earthquakes, planets, and
stars, are mentioned at least twice as much. (The
excellent Grade 2 FOSS unit and equipment kit,
Pebbles, Sand, and Silt, comes from California.
See Table 4.)
Canada (CMEC 1996) A Gr. 3 soil unit refers to soil components, the
interaction of soils with water, living things and
soils, and similarities and differences among soils.
A Gr. 10 Sustainability of Ecosystems unit includes
soil composition and fertility along with seven other
ecosystem concepts. (The “illustrative example”
focuses on soils.) Many provinces follow this
document, such as Ontario, the most populous
province (see below)
New York (NYSED 2009) Grades K-4 state that “soil is composed of broken-down
pieces of living and nonliving earth material.” Grades
5–8 refer to soil composition, soil monitoring, and
soil pollution. However, rocks and minerals, and the
moon, are all mentioned more frequently. Similarly
in Grades 9–12, where soil is mentioned briefly in
connection with ecosystems, while geological and
astronomical topics are mentioned more often
Ontario (Ontario MOE, 2007) In the Gr. 3 soil unit, students assess the environmental
impact of soils, and study the composition and
characteristics of different soils, and the relationship
between soils and living things. Detailed specific
expectations are given. Soil also appears as part of
the Gr. 9 Ecosystems unit, where students “plan and
conduct an investigation … into how a human
activity affects soil composition or soil fertility”
South Africa (DOE 2002, 2003; In Grades R-3, along with rocks, soil is mentioned, in
CAPS 2011) particular, the erosion of soil and the types of soil.
In Grades 4–6, the formation of soil, and the need
to maintain the fertility of soil, are mentioned in the
context of ecosystems, while the composition and
properties of soil are mentioned in the context of
earth changes
(continued)
Table 1 (continued)
Texas (TEA 2010) In Gr. 1, students sort components of soil by size,

texture, and color. In Grade 3, they study the
formation of soil by weathering of rock and the
decomposition of plant and animal remains. And in
Grade 4, they “examine properties of soils, including
color and texture, capacity to retain water, and
ability to support the growth of plants.” In Grades
9–12, soil is a significant part of one of the strands in
Environmental Systems
United Kingdom (UK DOE 2011) Key Stage 1 mentions several topics but not soil. Key
Stage 2 mentions some astronomy and environmen-
tal topics under physical processes and life processes,
but soil is not mentioned. In Key Stage 3, geological
topics such as rocks, and astronomical topics such as
earth and moon, are mentioned several times, but
soil is not mentioned. Similarly in Key Stage 4. It is
difficult to see “soil” as a content domain in this
science curriculum
United States (NAP 2012) Soil is one of the examples of “Crosscutting Practices.”
It is found in Life Sciences at each end level (Grades
2, 5, 8, and 12). It is frequently mentioned in Earth
and Space Sciences, in the subtopics of Earth
Materials and Systems, Plate Tectonics and
Large-Scale System Interactions, The Roles of Water
in Earth’s Surface Processes, Biogeology, Natural
Resources, and Human Impacts on Earth Systems
(Grades 2, 5, and 8)
Collins documents a declining trend of the number of undergraduate students

enrolled in soil science across the nation and notes that, based on anecdotal talks
with colleagues around the world, this trend appears to be international.
The frequency of soil science education in elementary and secondary is similar
to that at the university level. Table 1 summarizes references to soil in the curricu-
lum documents of various English-speaking countries and regions. Soil science is
explicitly mentioned in Canada, South Africa, the United States, and their provinces
and states, although other earth and space topics such as geology and astronomy
appear more frequently. In contrast, soil science doesn’t appear as a unique content
topic at all in most European national curriculum documents, although, as mentioned
above, soil educators in Europe are active in promoting soil education to adults and
children in events outside of the classroom (i.e., Blum and Kvarda 2006; Broll 2006,
2009; Creamer 2009; Hallett 2009; Houskova 2009; Towers et al. 2010). In Africa,
a region where many families live close to the land, information is limited and what
is available provides a mixed picture. Soil science doesn’t seem to feature promi-
nently in the Nigerian science curriculum (Oludipe 2011). In parts of Ethiopia,
however, soil fertility and water and soil issues are an important part of the work of
school environmental clubs, where students implement environmental conservation
at school compounds and family lands (Edwards et al. 2010).
6 D. Hayhoe
This article reviews existing research on soil science in elementary and secondary
teaching and learning, and summarizes the English-language resources available to
increase understanding of soils and awareness of the essential role they play in sustain-
ing life on Earth.
2 Results of Research on Elementary and Secondary

Teaching and Learning About Soils
In this section, studies on student and teacher initial understanding about soil, and
what programs have been effective in helping them understand soil better, are
grouped by level (elementary or secondary) and by subject (students or teachers).
2.1 Elementary Students’ Understanding of Soil
2.1.1 Initial Understandings
This section talks about four studies that report on elementary students’ initial
understanding of soil concepts (Table 2). Several other studies report pre-post gains
in understanding of soil concepts (Table 3), but did not report on the student’s initial
understandings as demonstrated on the pre-tests.
Geyer et al. (2003, 2004) worked with 150 children, 4–11 years old, in primary
schools in Germany. The purpose of the study was to identify aspects of soil and
agricultural ecology that could be taught to children at different ages, whether the
children could understand the three-dimensionality of soil and its interactions with
ecosystems, and how soil science learning works with very young children. The
children were introduced to the program with the question: “Why should soil be
interesting for you?” The youngest children referred to soil as a playing-ground,
while the older children had some idea of soil’s three-dimensionality, and referred
to seeds, plants, trees, and earthworms, which all live in the soil (Fig. 2).
The researchers also asked the 150 children to draw pictures that illustrated their
ideas about soil. These pre-intervention pictures revealed the following initial
understandings of soil:
• 4–7 years: children set the horizon (the ground) at the very bottom of the picture.
They don’t have any place in the diagram or in their mind for ‘soil’
• 7–9 years: some drawings allow for space for soil, but this is not developed
• 9–11 years: they already have an idea of how soil could look, its genetic pro-
cesses, and to some extent, it’s physical properties
Happs (1981, 1984) interviewed 40 students in Waikato, New Zealand from
Gr. 7 to university level. The study was concerned about what students think of the
nature, origin, age, and depth of soil, along with changes that might occur in soil.
Table 2 Initial understandings of soil by elementary students in various countries
Reference Country Students Probe used Topics studied Initial student understandings
Geyer et al. 2003, Germany Age 4–11 Interviews and Purpose of soil; 2 or 3D Age 4–7 no idea of anything below
2004, 2006 labeled drawings picture of soil ground
Age 7–9 some idea of depth, but not
soil
Age 9–11 have some pre-ideas of soil
Happs 1981, 1984 New Zealand 40 Gr. 7 students; Interviews; Makeup, origin, depth, and The majority of students thought that
(221 Gr. 7–12 (followed by a age of soils. Changes that soil (1) is a medium for plant
students) Likert scale test soils undergo growth and a home for small
survey of 16 animals, (2) has always been there
items) (a few said that it formed from
organic matter), (3) is as old as
the Earth, (4) is between a few cm
and 10 m deep (a few said that it
was several thousand km deep)
and (5) does not change, or,
Surprising Facts About Soils, Students and Teachers…
changes to clay and then to rock

in some undefined cycle. Many of
the students used the terms “dirt”
and “soil” synonymously
Russell et al. 1993 UK 58 Gr. K-6 students Student log books, Makeup, origin, and After exploration activities, to
(5–11 years) drawings, permanence of soil. familiarize them with soil, but
discussion. Changes to soil that might before intervention strategies,
What is soil? occur and their reasons students thought the following:
What is in it? Soil’s use or function: for growing
plants
Age 5–8: soil is mud, sand, or stones
Age 8–11: some idea of sand as a
mixture
Little idea of changes to soil, or its
7
origin
8
Table 3 Pre-post gains in soil understanding by elementary students in various countries

Reference Country Students Probe used Topics studied Treatment Results
Geyer et al. 2003, Germany Age 4–11 year Interviews and Purpose of soil, ideas teacher lessons, soil After some weeks,
2004 labeled diagrams about soil layers, animals, fieldwork in diagrams
soil and ecosystems a pit, observations of included a lot
a variety of soils more detail about
soil layers
Gulay et al. 2010 Turkey Preschool children 12 test items Characteristics of soil, 9-day program in four The experimental
(5–6 years), administered living beings different nursery group achieved
from low orally with figures on/under the soil, classes with significantly
socio-econ. and photograph, importance of soil, activities on higher than the
back-ground, pre-test, post-test, reasons and results characteristics of control group on
with little and delayed of erosion soil, living beings on the post-test,
knowledge of post-test or under the soil, although both
environ. subjects importance of soil, groups were
protection of the soil, similar on the
and identification of pre-test
reasons and results
of erosion
Lippert 2006 US 97 Gr. 7 students 26 multiple-choice A variety of mostly Web-based module for Significant gains
pre-post test items factual questions 2–3 days, based on were found on
related to soil slides used in most of the items
D. Hayhoe
extension education
Reference Country Students Probe used Topics studied Treatment Results
Randler and Germany 123 Gr. 5–6 students Pre-post tests with Three experiments: Teacher-centred No significant
Hulde 2007 5 open-ended Water-holding presentation of difference in
questions related capacity of the moss, experiments versus pre-post gains for
to content erosion of grassland learner-centred both groups for
versus agricultural hands-on work post-tests held
land, water-cleaning immediately after
capacity of soil treatment. For
post-tests held a
month later, the
learner-centred
group scored
significantly
higher
Russell et al. 1993 UK 58 Gr. K-6 students Student logbooks, Makeup, origin, and Detailed intervention Better knowledge
(5–11 years) drawings, permanence of soil. strategies over that soil is made
individual Depth of soil. 5 weeks, involving a of living things,
discussion, and Changes to soil that variety of strategies. has particles of
pre-post might occur and different size, and
intervention their reasons owes its origin to

interviews the transforma-
tion of previous
inorganic
substances
9
10 D. Hayhoe
Fig. 2 A mind-map of children’s previous concepts of soil (Geyer et al. 2004)
Students were first given a variety of familiar materials related to soil (loose portion
of topsoil, section of turf, grass with root system, clay, sawdust, potting-mix, pebbles),
and asked to identify what they saw, as a help in eliciting from them their concept
of soil and soil development. Various stimulus words were also placed on cards to
further aid the questioning: soil, colour, silt, rocks, sand, clay, consistence, texture,
structure, profile, living things, vegetation, water, parent material, etc. A further
multiple-choice survey was constructed on the basis of the interviews and adminis-
tered to an additional 221 middle and secondary school students.
Although nearly everyone described soil as providing support for life, many also
referred to it as “dirt.” Almost half thought that soil was formed the same time as the
earth, i.e., having an age as old as 100,000,000 years, although a few thought it might
be only 20 years old. Some saw soil as the product of rotting vegetables and animals,
a few as originating in volcanic ash. Some upper school and university students rec-
ognized that soil development was a “multi-source mechanism.” Most recognized that
soils continually changed over time, but couldn’t describe how. Only in their estimate
of the depth of soil were student answers close to the scientific view: one third thought
that soil was under 1 m in depth, and another third between 1 and 10 m in depth.
Russell et al. (1993) worked with 58 children between 5 and 11 years of age.
They first engaged the students with exploration activities such as looking through
soils with a magnifier, thinking about which soils would be best for plant growth,
and looking down a hole outside to think about how deep soil might go if you could
dig as deep as you wanted. They followed this exploration stage with pre-intervention
interviews to elicit the students’ views on soil. These interviews focused on the
following topics:
• The function of soil: More than half of students didn’t know or had no response.
About a third thought that soil was for growing plants.
• The nature of soil: Almost half of the students made no reference to soil compo-
sition; 12 students (mostly upper-level) referred to soil as a mixture; 8 of the
youngest students referred to soil as mud or sand.
• Changes in the properties of soil: About a third of the students mentioned changes
in water content or wetness of soil.
• The origin of soil: Few students offered ideas as to where soil came from. Some
referred to the formation of soil over millions of years, from rotting vegetation,
as well as sand or gravel.
The students were also asked to make detailed drawings of what they thought
was under the ground. They may have been given more direction than for the draw-
ings reported above by Geyer et al. (2003, 2004), as the results were consistent but
also more specific:
• Younger children tended to draw ‘no-layer’ diagrams
• Other students drew various layers under the ground, including such things as
soil, clay, sand, lava, or where they found worms, pipes, bones, tar, stones, rats,
and Earth’s core
• Some of the older students clearly marked out a layer of soil in their drawings
(Fig. 3)
The research summarized by these four articles was extensive in nature, and
covered a span of ages from 5 to 12 years. It included student discussions and inter-
views as well as analysis of labeled diagrams and pictures drawn by students.
Similar results regarding student views of the nature and function of soil, changes in
soil, and the origin and age of soil emerge. Before instruction, students of all ages:
1. Are unclear about the nature and composition of soil
2. Have some idea of a layer of soil under the ground
3. Tend to think of soil as supporting plant growth and as a home for other forms
of life
4. Have little idea of the age of soil, often considering it to be millions of years old
5. Have little idea of how soil forms from weathering and erosion, which contrib-
utes to its composition of sand, silt, clay, and humus
6. Have some idea of the depth of soil, closer to reality than for the other attributes.
Several qualifications should be noted about this work. First, the students
involved in the research appeared to be all from urban schools. Students growing up
in rural communities, especially those living on farms, may have given more
informed answers. Second, the students in these three reports were from similar
Western European cultures. Would students from urban schools from other cultures
have responded differently? Third, if this same research had been done with
students in North American provinces and states where soil had already been
studied as a formal topic in primary school, such as California, Ontario, or Texas
(see Table 1), would the data from junior students be substantially different than
what we saw here? I am not aware of any data on this question, since soil is not
usually one of the topics included in international science assessments. Fourth,
12 D. Hayhoe
Fig. 3 Initial drawing, before treatment, by a junior level student in England, showing an unusual
number of references to minerals (Russell et al. 1993)
these three studies were conducted 10, 20 and 30 years ago. When the astronomical
beliefs of Gr. 6 British students were surveyed in the mid 1990s (Sharp 1996), for
example, it was found that the students knew significantly more about astronomy
than Gr. 6 British students who had been surveyed only a decade or so earlier (Baxter
1989). This advance was attributed to the great increase in number and quality of
astronomy and space programs appearing on television in the intervening decade.
Might the same be true of soil science over the past two or three decades? The U.S.
Smithsonian Soils exhibit was seen by millions in its 18-month showing in 2008–
2009 (Collins 2008; Megonigal et al. 2010). The more permanent “Underground
Adventure” soil exhibit in Chicago’s Field Museum has also been seen by many.
There is also an increasing amount of high quality soil education outreach taking
place across Europe, coordinated by the European Network of Soil Awareness
(Broll 2011), as well as soil education websites in many countries (see below).
In general, however, there is no indication of a widespread change in soil science
information available to students and the most likely assumption is that initial
understandings of most students today have not changed.
2.1.2 The Effect of Treatment Activities
Five studies have attempted to quantify the effectiveness of various “treatment” or

“intervention” programs for increasing elementary students’ understanding of soil
concepts to inform teaching strategies over a range of ages (Table 3). Gulay et al.
(2010) assessed Turkish 5 and 6 year old’s understanding of five aspects of soil: its
characteristics, living beings that live in or under it, its importance, protection of the
soil, and the causes and effects of soil erosion. The subjects were divided into control
and experiment groups, both of which received the pre-test, post-tests, and delayed
post-test (2 weeks later), but only the experimental group was exposed to the 9-day
program of treatment activities. The children were selected on the basis of two
criteria: (1) a low socio-economic background, and (2) not having been exposed to
any prior education on soil, erosion, and the environment. Treatment activities
included story, games, drama, songs, fieldtrip, experiment, art, and work with soil in
a corner of the classroom. The program focused on a puppet, Tipitop, and was called
“We are Learning about the Soil with Tipitop and His Friends.” The pre-post tests
consisted of 12 questions administered orally by adults along with slide figures and
photographs. There was no significant difference in pre-test scores between the
experimental and control groups. The experimental group performed significantly
higher on post-tests scores and delayed post-test scores than either the control group
or their own pre-test scores. Unfortunately, no details were given as to which soil
concepts were best understood initially or which were learned most effectively dur-
ing treatment.
Geyer et al. (2003, 2004) also worked with children from 4 to 11 years in age.
Teacher lessons and fieldwork involved students examining soil pits, various soil
layers, and animals that live in the soil. Interviews took place before and after the
fieldwork, and students submitted labeled diagrams before, immediately after, several
weeks after, and a year after the classes on soils. Diagrams submitted after fieldwork
included more realistic colours and identified layers of the excavated soils. Students
remembered soil layers weeks and months later. Figure 4 demonstrates one 9-year-old
student’s diagrams before the fieldwork, immediately after, several weeks later, and
a year later. Before the lessons, soil is merely a playing ground. After fieldwork, the
student documents the layers. 6 weeks later, soil has become part of his life, and a
year later, he still remembers the layers of soil. It is now part of his ‘normal life.’
After analyzing hundreds of diagrams from the 150 students aged 4–11 years, drawn
before and after the fieldwork lessons, the authors concluded that:
• Students aged 4–7 years could remember soil layers, after fieldwork, but were par-
ticularly interested in soil animals, which they “drew frequently and very exactly”
• Students aged 7–8 years remember soil colours and soil genetic processes
• Students above 9 years in age are interested in the science and ecology of soil,
and remember genetic processes and some physical properties very well. Animals
and plants are less important. They understood the difference in forest ecosys-
tems and agriculturally managed ecosystems. They were also able to make their
own conclusions to their studies.
14 D. Hayhoe
Fig. 4 Diagrams made by a 9-year-old student before the soil fieldwork (left), soon after, 6 weeks
later, and a year later, showing progress and permanence in learning soil layers (Geyer et al. 2004).
Notice the surprising amount of underground soil features in the layers drawn in the last diagram
Russell et al. (1993) also involved students aged 5–11 years. Following initial
interviews, students were exposed to intervention strategies (i.e., classroom
activities) over 5 weeks. The intervention phase of Russell et al. employed a
number of strategies, including encouraging the children to evaluate their ideas
side-by-side with the “right” ideas, develop more specific definitions for soil-
related words, generalize across concepts, and use secondary sources of informa-
tion. The specific soil intervention activities consisted of teaching the children to
take a closer look at soil, compare different soils, and develop ideas about what
is under the ground.
Several weeks later, a set of data was elicited from the students complimentary
to the pre-test data (log books, drawings, discussions, post-intervention interviews)
and was analyzed to reveal:
• The nature of soil: Younger children (age 7–9) initially had difficulty understanding
that soil was composed of various materials such as sand, clay, and living mate-
rial. When stones were rubbed together, producing dust, the children still had no
idea that this had anything to do with the origin of soil. When they started strain-
ing soil through cloth material, however, they began to see that it was made up of
several different components. Eventually they were able to conceptualize soil as
made up of several organic and inorganic components, although none of them
mentioned air as one of these.
• Comparing soils: Although some children tended to judge the ‘goodness’ of soil
at first by appearance only, they soon learned to use better criteria (i.e., a fair
test), including how well the soil supported plant growth.
• Thinking about what’s under the ground: While a few hands-on activities related
to this topic (i.e., digging a hole in the garden 1 m deep), little time was left to
consider secondary sources. Many children, however, began to understand that
underneath soil we come into contact with rocks. (Rocks were the next activity
studied in this intervention phase, after soils.)
Post-intervention interviews revealed a complex picture of both learning and
un-learning that took place concerning the nature of soils and what is in soil,
including:
• Living things: More students referred to living things as one of the constituents
of soil after intervention than before (57% vs. 27%), where living things could be
plants, roots, seeds, microorganisms, or small creatures.
• Soil constituents: While younger students mentioned fewer constituents of soil
after intervention than before, perhaps because they un-learned several things
that they previously thought were constituents, older students mentioned more
constituents after than before!
• Organic matter (dead not living): The same number of students mentioned
organic matter after intervention as before (61% vs. 60%).
• Inorganic matter: After intervention approximately the same number of students
mentioned
• Particle size: more students after intervention referred to different-sized particles
in soil than before intervention (74% vs. 31%).
• Origins of soil: Before intervention, 28 students referred to the translocation of
soil from another location, while eight students referred to both translocation and
transformation. Only eight students mentioned soil being transformed. After
intervention, 27 students referred to the transformation of soil, while only 18
referred to its translocation. (Often, students referring to the translocation of soil
thought of it as coming from gardens, garden centres, etc., brought by humans.)
• Nature of soil transformation: More students thought of soil as having inorganic
origins after intervention than before (26% vs. 12%), some mentioning volcanoes
sending forth lava, rocks being ground down, sand coming from the sea, things
colliding together, etc.
• Types of soil: When students were shown five samples of soil, and asked to
classify each as soil or non-soil, pre-post results were mixed concerning three of
the samples, showing the continued subtlety of understanding what soil really is,
given various human perspectives of soil. (The three samples included sandy
topsoil, chalky soil, and damp peat.)
16 D. Hayhoe
Lippert (2006) used a web-based basic soil module with pop-up test and
audio files as a treatment for 97 Gr. 7 students in South Carolina. Students
answered a multiple-choice pre-test in class, then studied the web-based module
over 2–3 days in a computer lab, and then answered the post-test in class.
A month later, they answered the same post-test again. The purpose of the
research was to see whether a web-based module was effective in instructing
students on soils. Results were positive, with pre-post gains on 21 of the ques-
tions exceeding 30%. Three questions showed moderate gains (20–29%), and
two questions showed little gain (10–19%). When the same module and tests
were given to 150 university students, results were only slightly better. Results
on the delayed post-test indicated a drop-off in knowledge from the post-test
scores taken immediately after the module, although there were still significant
gains over the pre-test (Table 4).
Insufficient information is available to draw conclusions about the gains for
specific items. In addition, the gains made by the students for many of the 26
questions are quite significant, after only 2 or 3 days (periods?) in the school’s
computer lab. This contrasts with the 5 weeks of intensive interventionist strate-
gies employed by teachers in the study with Kindergarten to Grade 6 study
reported by Russell et al. (1993), where the gains did not appear to be as significant.
Finally, most of these 26 questions appear to be concerned with rather factual,
technical details, and not concepts that lie at the heart of understanding soil
science, which can be deeply engrained in student’s thinking through hands-on
investigations and minds-on discussion and questioning. The fact that the
delayed gain (a month later) was much lower than the immediate gain or many of
the questions appears to bear this out.
In another German study reported on by Randler and Hulde (2007), 123 students
enrolled in two Gr. 5 and two Gr. 6 classes in a German middle school were given a
pre-post test of five open-ended questions, with an intervening treatment program
consisted of three different ecological experiments dealing with soil ecology: (1)
investigating the water holding capacity of moss, (2) studying the erosion of grass-
land versus agricultural land, and (3) finding the water cleaning capacity of soil. The
principal variable tested was the effect of learner-centered vs. teacher-centered
classrooms. One Gr. 5 and one Gr. 6 class received the learner-centered treatment,
with the other two receiving the teacher-centered treatment. In the teacher-centered
classes, the teacher carried out the experiments and discussed the results with the
students; in the learner-centered classes, the students carried out the experiments.
The pre-tests were completed just prior to the teaching, the first post-test was car-
ried out just after the teaching, and a delayed post-test was carried out4 weeks later.
For the post-tests, two additional questions were added to the same five used in the
pre-test. The following are example items of the pre-post tests:
• Which specific characteristic is especially related to moss? (water holding
capacity)
• What specific material from everyday life has a similar characteristic? (sponge)
Table 4 Gains on 26 knowledge questions about soils (Lippert 2006). Grade 7 students studied a
web-based module on soils for 2–3 days, and answered a pre-post test with the following question
stems
Question stem Pre-post gain Delayed gain
Soil is roughly what percent pore space? 56 35
The three particle sizes for soil minerals do not include: 10 8
The smallest soil particle is: 62 53
A texture triangle tells us: 31 22
Which statement is true? 77 67
Clays generally have: 69 52
Horizons: 11 5
The soil horizon which loses minerals and clay to the 36 16
layer underneath it is labeled with the letter:
Bedrock breaks up because of: 26 12
Undeveloped soils have: 50 43
The five soil forming factors are climate, topography, 11 5
biology, time and:
A soil will develop the fastest when the weather is: 49 34
Topography refers to: 33 26
Most organic matter is decomposed by: 23 21
In general, it takes about how long to form a layer of 51 44
soil the thickness of a sheet of paper?
For plants to grow, they need how many nutrients? 50 40
Secondary plant nutrients are: 20 27
Which is correct? 38 34
If phosphorus is deficient in the soil, the plant leaves 70 38
appear:
When a plant is deficient in potassium, the leaves: 36 20
Phosphorus doesn’t move through the soil with rainfall 35 13
because:
When a positively charged atom takes the place of 43 43
another positively charged atom on clay, it is called:
An acid soil: 39 19
Soil acidity is not formed from: 39 15
For maximum plant nutrient availability, the ideal soil 45 28
pH should be close to:
Erosion always occurs when there is: 42 32
• Water above the ground is often dirty, ground water is nearly clear. Explain.
(Plants and soil material filter dirty water.)
• Steep sloes often are planed with grass. What is the advantage? (Protects soil
from erosion)
• What would happen if the soil is bare (without plants) (erosion would take away
the soil)
18 D. Hayhoe
Table 5 Significant gains retained after a 1-month delay (Randler and Hulde 2007). Grade 5–6
students studied the water holding property of moss, soil erosion on grasslands compared with
agricultural lands, and the water cleansing capacity of soil, and were tested for pre-post gains in
understanding, in teacher-centered versus learner-centered environments
Test Treatment Mean Out of SD t-value Probability
Pre-test Learner-centered 1.37 5 0.89 −0.227 0.821
Pre-test Teacher-centered 1.40 5 1.00
Post-test Learner-centered 5.50 7 0.91 −0.588 0.557
Post-test Teacher-centered 5.60 7 1.08
Delayed Learner-centered 5.38 7 0.94 2.579 0.011
Delayed Teacher-centered 4.91 7 1.06
Average scores were statistically the same for both treatment groups (Table 5).
For the delayed post-test, however, the mean score of the teacher-centered groups
had declined somewhat from the post-test scores immediately after the teaching,
whereas the mean score of the learner-centered group remained the same, resulting
in a significant difference between the two groups. The main conclusion from this
is straightforward: learner-centered classrooms, where students engage in hands-on
experiments, rather than just watching and listening to the teacher doing and dis-
cussing the experiments, results in significantly better long-term retention of the
learning. This conclusion is consistent with a wide body of research concerning
student learning in science.
2.1.3 Summary
Elementary students are initially unclear regarding the nature, composition, and
function of soil is concerned, although they usually know that it supports plant
growth in some way. They have little idea of the origin, age or formation of soil,
although many of them, especially middle school students, understand that soil is
not that deep and is often sitting on top of rock (Happs 1984; Russell et al. 1993).
Nevertheless, effective education programs can be mounted with students of all age
groups, from pre-school to middle school. While a short-term teacher-centered pro-
gram may lead to strong initial gains in student understanding of some facts and
details about soil (Lippert 2006), an extensive 4–5 week program of hands-on explo-
rations combined with “minds-in” discussions for elementary students is likely
required to permanently alter student understanding of key soil concepts (Geyer
et al. 2003, 2004; Russell et al. 1993; Randler and Hulde 2007).
2.2 Elementary Teachers’ Understanding of Soils
Elementary teachers’ understanding about soil may be just as important as that of

students, especially in countries where soil is taught as a classroom subject. Very
little research has been reported on this subject, however.
2.2.1 Initial Understandings
In a study of 108 elementary and middle school teachers in Nebraska, only one of
the 38 items concerned soils Gosselin and Macklem-Hurst (2002), and this item
elicited the lowest score of the test. While average scores on the 38-item pre-test
were 55%, only 16% of students correctly disagreed with the statement, “Soils are
deposited as natural rock layers.”
In another study on 87 preservice elementary teachers in New York State, a pre-test
was administered that included writing the definition of clay, listing products made from
clay, and explaining the origin of clay (Rule 2007). Only a minority of the preservice
teachers thought of clay as a natural substance in the Earth (Table 6). Most naturally
thought of modeling clay without making any connection between this and the Earth’s
natural substances. As far as the origin of clay was concerned, only three of the eleven
suggestions could be considered scientific: that clay forms in the ground (nine responses),
that it forms from chemical weathering (5), and that clay minerals are found in soil (4).
Since soil science is a topic at the Grade 3 level in Ontario, a study was con-
ducted on preservice elementary teachers’ initial understandings of a complete set
of soil concepts (Hayhoe et al. in-press). Seventy four primary-junior (K-6) teachers
out of a potential pool of 125, studying at a medium-sized public university near
Toronto, voluntarily responded to a 32-item multiple-choice questionnaire. The pre-
service teachers represented the cultural diversity of the greater Toronto area, and
all had a university degree, but very few in science. The teachers achieved a mean
of 55% on the questionnaire, in contrast to a random score of 25% (Table 7).
In a further study, 25 of the 32 items were given to a second group of 98 preservice
elementary teachers at the same institution, and to 41 preservice elementary teachers at
a local private Christian university. The results were all very similar (Hayhoe et al.
Manuscript submitted for publication). The results in Table 7 indicate that while these
Canadian teachers understand a significant number of important soil concepts, they
have misconceptions or lack of knowledge on many others and most likely need some
instruction on soil to be able to successfully teach it to Grade 3 students. (It is important
to note that most of these Ontario teachers had not received soil science education in
their own Grade 3 schooling; it did not enter the curriculum until 1999–2000.)
2.2.2 The Effect of Treatment Activities
When geoscience topics such as soils are included in science methods courses taken
by preservice elementary teachers, is there a significant gain in their pre-post test
scores? In Gosselin and Macklem-Hurst (2002), students met twice a week for a
total of 4.5 h. The course content “was primarily presented through the use of both
hands-on and minds-on approaches, including inquiry-based activities.” Two col-
laborative projects concluded the course, one concerning weather phenomena, and
the other concerning stream flow data. Post-test scores were then collected. The pre-
post gain on the single item about soils was 16% (16–32%), which compares unfa-
vorably with the average on the 38 items of 25% (55–80%). After instruction, most
students still thought that soils were deposited as natural rock layers.
20 D. Hayhoe
Table 6 Nature and origin of clay as understood by 87 preservice elementary teachers in New
York State (Rule 2007). The pre-test included writing the definition of clay, listing products made
from clay, and explaining the origin of clay
General category of definition of soils No. of teachers
Unspecified moldable substance, used to make things, to be shaped 39
Natural substance found in the Earth, sediment found along riverbanks 39
Raw material for ceramics, used for pottery, solid substance 27
Art material, easy to form, used for making things 20
Manufactured artificial material, play dough 18
Material for making models, modeling clay 17
Type of soil, hard soil, thick soil, soil in the ground 13
Wet dirt, mucky substance reddish brown in colour, mud 11
Category of concept about origin of clay
Formed by a mixture of particles with water. One of the layers within rock. 18
Made from the breakdown of rock, a result of heating of rock, etc.
Pressure is needed for clay mineral formation. Pressure on soil and water plus 12
time produces clay. It forms over years from compression and mixture of
minerals and water
Clay minerals are ground rock. Clay comes from the breakup of larger 12
minerals into clay
Clay forms from a mixture of materials – sediments and minerals 9
Clay forms in the ground, from a specific mixture of minerals in the ground 9
Heat, melting is involved in clay formation; melting caused by the heat of the 9
Earth
Clay was here from the beginning of the Earth. God made it 7
Chemical weathering forms clay; it originates from a series of reactions at 5
Earth’s surface
Clay minerals are found in soil. Minerals come from the soil 4
Clay minerals originate from organic materials. The minerals come from 4
plants
Evaporation and recycling forms clay 3
In a follow-up study by Hayhoe et al. (Manuscript submitted for publication), 19

teachers at the private Christian university participated in an in-class treatment con-
sisting of two 3-h classes of hands-on activities and discussion related to soils,
together with some after-class work (Hayhoe et al. 2011). The class activities
involved up to ten hands-on experiments – ones that students studying the Grade 3
soils unit would typically do over a period of several weeks – together with group
and class discussions and readings (Fig. 5). Five months after the activities on soils,
and 4 months after the course was over, the same post-test questionnaire was given
to the teachers so that long-term pre-post gains on understanding of soil concepts
could be analyzed.
An “environmental attitudes” survey was also developed and administered simul-
taneously with the soil concepts questionnaire. The preservice teachers answered a
20-item Likert scale survey to test their attitudes towards five environmental topics:
climate change, energy usage, water, nuclear energy, and soils. The researchers
Table 7 Pre-test selections of 74 preservice teachers on soils (Hayhoe et al., manuscript submitted for publication). Preservice elementary teachers at a
mid-sized public university in Canada were tested for initial understanding of different soil concepts
Some correct responses selected by more than two-thirds of the teachers %
Sand is mainly formed on Earth by weathering of rocks 89
Earthworms mix the soil and allow for more air and water to be in it 88
When trees and plants die they decompose into soil nutrients used for plants to grow 84
Soil is essential for life and society to survey 84
Compost improves plant growth … by adding essential nutrients to soil 77
The component of soil that is slippery or sticky and keeps its shape after you let it go is clay 74
Rodents do not function as soil decomposers 73
Crop rotation helps maintain fertile soils 73
The life forms that live in the soil are decomposers 64
Some correct responses selected by less than one half of teachers % Some incorrect responses selected %
One handful of fertile soil is likely to have more organisms than people living on earth 14 … people living in a small village 38
… people living in a small city 45
It usually takes 100–1,000 years to form 1 cm of topsoil 20 … 1–10 years to form 1 cm of topsoil 66
When we look at soil with a magnifier, we are very unlikely to see soil fungi 27 … plant roots 28
… silt 35
Soil is composed of solid particles with spaces between them for air and water to enter. In good 27 … 25% 58
soil, approximately 50% of the total soil volume is space for air and water
A very fine component of soil that feels like powder when it is dry is silt 45 … sand 28
… clay 23
Clay particles are the smallest particles in soil 7 Silt particles … 46
Sand particles … 34
You place equal amounts of sand, clay, and silt in three different test tubes of water, shake the 38 … with clay … 45
test tubes, and sit them in a stand. In the test tube with sand, the solid will settle down first.
Decayed organic matter in soil is called humus 26 … decomposers 45
The role of humus in soil is to provide nutrients for micro-organisms and plant roots 28 … is to retain water for dry periods 41
One of the functions of soil is filtering impurities out of water 43 … supplying oxygen for us to breathe 22
21
… none of the above 30

22 D. Hayhoe
Fig. 5 Preservice elementary teachers studying soil observe its different components, notice its
formation from the erosion of rocks, and ponder the role earthworms in it (Hayhoe et al. 2011)
wanted to see if concern for soils was correlated with initial understanding of soil
concepts, and if exposure to soils activities increased concern for soils (in contrast
to the other four environmental topics) as well as understanding of soil concepts.
This 32-item soil questionnaire and 20-item environmental survey were given both
to the 19 students from the private Christian university (Tyndale), in pre-post posi-
tions, as well as to the 74 students from the mid-sized public university mentioned
earlier (UOIT), in pre-test position only. (Only 67 of the 74 students from the mid-
sized public university completed both the soil questionnaire and the environmental
survey.)
For the pre-test questionnaire and survey, the means for the two universities were
the same, 57% for the soil concepts 32-item MC test and 75–77% for the 20-item
Likert scale environmental survey, which measured their attitudes toward soils
(Fig. 6). For the post-test surveys, the means for the 19 teachers from the smaller
university went from 57% to 78% for the soil concepts and from 75% to 88% for the
soil attitudes (Fig. 7). This research indicated that 10% of class time in a science
methods course can significantly affect both the environmental concern and the con-
ceptual understanding of teachers for soils, although an increase in the one was not
correlated with an increase in the other (i.e., teachers who increased the most in
their environmental concern for soils did not necessarily increase the most in their
understanding of important soil science concepts).
In a second year of this study, the 32-item soil survey was reduced to 25 items,
by removing items on which teachers had initially achieved 90% or so on previous
pre-tests, and a few with poor discrimination indices (Hayhoe et al., Manuscript
submitted for publication). When this 25-item soil concept survey was applied to a
new cohort of preservice teachers at both the private and public universities, and the
previous years’ results were re-analyzed, the 2 years of data were very similar
(Table 8).
The consistency of these results suggests that any effect is not peculiar to one
particular cohort or university. The only significant effect was the pre-post gain:
teachers gained 37% ([67.1−48.1]/[100−48.1]) of what was lacking in their under-
Fig. 6 Teacher 75% 77%

understanding of soil 80%
concepts and attitudes toward 70%
soils (Hayhoe et al. 2011). 57% 57%
Preservice elementary 60%
teachers at a small private 50%
Christian university in Tyndale 19
Canada (Tyndale) performed 40% Pretest
the same as their counterparts 30%
at a mid-sized public UOIT 67
university (UOIT) on pretests 20% Pretest
for soil concepts and soil 10%
attitudes
0%
Soil Soil
Concepts Attitudes
Fig. 7 Pre-post gains in 88%

teacher understanding and 90%
attitudes towards soil 80% 75%
(Hayhoe et al. 2011). 70%
Preservice elementary 70%
teachers at a small private 57%
60%
Christian university in
Canada (Tyndale) gained in 50% Tyndale 19
both soil concepts and soil Pretest
40%
attitudes, although there was
no significant correlation 30% Tyndale 19
between gains for individual Posttest
20%
teachers
10%
0%
Soil Soil
Concepts Attitudes
standing of these soil concepts. Comparing Table 9 with Table 7, on some items the
preservice teachers made large gains: understanding how many life forms are in a
handful of soil, how many years it takes for soil to form, what decayed organic
matter is called (humus) and what it does, which component of soil settles down
first in water (sand), and how to differentiate between dry soil and dry clay by tex-
ture. On some other items, they made modest but significant gains – 50% of good
soil is space for air and water, the smallest particles in soil are clay particles, soil
fungi are too tiny to be seen with a magnifier, and soil filters impurities out of our
water – although on the delayed post-test scores, still only a minority of the preser-
vice teachers answered these questions correctly.
24 D. Hayhoe
Table 8 Pre-test scores and gains on 25 items related to specific soil

concepts, by preservice elementary teachers at two universities over
2 years (Hayhoe et al., manuscript submitted for publication)
Application of the
soil concept test Participants Raw mean st % mean
Year 1
UOIT pre-test 74 12.53 3.96 50.1a
Tyndale pre-test 21 12.43 2.53 49.7a, b
Tyndale post-test 21 16.57 3.61 66.3a, b
Year 2
UOIT pre-test 98 12.45 4.23 49.8a
Tyndale pre-test 20 12.00 3.74 48.0a, b
Tyndale post-test 20 16.95 4.41 67.8a, b
a
The differences between the four pre-tests (two institutions over 2 years)
were not significant
b
The differences between pre-post tests for both years were significant
(p < .001)
Table 9 Pre-post mean item scores averaged over 2 years for selected soils items with 41 preser-
vice elementary teachers (Hayhoe et al, manuscript submitted for publication). The pre-test means
in this table are for the 41 Tyndale preservice teachers, whereas the pre-test means in Table 7 are
for 74 UOIT preservice teachers
Pre-test % Post-test
Pre-post test scores for items showing significant gains. Correct Mean (%) mean (%)
answers are given with the item stems below (n = 41) (n = 41)
One handful of fertile soil is likely to have more organisms than 12 71
people living on earth
It usually takes 100–1,000 years to form 1 cm of topsoil 10 60
When we look at soil with a magnifier, what are we very unlikely to 31 43
see? Soil fungi
Soil is composed of solid particles with spaces between them for air 26 36
and water to enter. In good soil, approximately how much of the
total soil volume is space for air and water? 50%
A very find component of soil that feels like power when it is dry 36 62
is silt
What are the smallest particles in soil? Clay 10 24
You place equal amounts of sand, clay, and silt in three different test 43 67
tubes of water, shake the test tubes, and sit them in a stand. In
which test tube will the solid settle down first? Sand
Decayed organic matter present in soil is called humus 12 48
The role of humus in soil is to provide nutrients for micro-organisms 45 62
and plant roots
Which of the following functions does soil do? Filter impurities out 24 31
of our water
2.2.3 Summary
Of the three studies on elementary teacher understanding of soils, the first only dealt
with one concept (soil not being deposited like rock layers). The second only dealt
with concepts related to clay, although in great depth. Only the third set of studies
reported on a variety of soil science concepts. These found that preservice elemen-
tary teachers, most of whom had not studied science at the post-secondary level and
probably never studied soil science, nevertheless initially understood many impor-
tant concepts about soil. At the same time, they had many misconceptions or areas
of ignorance about soils. Long-term gains made after only 2–3 classes in their sci-
ence methods course reduced by a third what was lacking in their knowledge and
understanding of soil science concepts. This is probably all that can be expected in
a treatment program covering many soil science concepts in a modest amount of
class time and readings.
2.3 Secondary School Students’ Understanding of Soils
Only two studies report on secondary school students’ initial understandings of

soil (Table 10). The findings of Happs (1981, 1984) have already been reported
(Sect. 2.1.1). Drieling (2006, 2008) examined the ideas of senior students in
Germany, 15–16 years of age, through the use of interviews and drawings. One
student imagined that below the ground there was uniform matter where animals
live and which perhaps had water underneath. Another students envisioned definite
soil layers below the ground “in the sense of divided geological layers of different
materials such as soil, sand, gravel, or rocks.” A third student remembers digging
a hole in the woods and seeing many different layers of colours and compositions
in the soil below the ground. Drieling concluded that there was a wide variation
among the students as to how closely their ideas approached the scientific under-
standing about soil layers and profiles.
Three studies have reported on effective treatment activities on soil under-
standing at the secondary level (Table 11). Cattle et al. (1995) introduced the use
of computers in teaching detailed processes of soil science to upper secondary
school and university students. Since technology has developed greatly over the
intervening years, the details of this program are probably not of much use now.
Drieling (2008) proposed using a constructivist model of activities for working
with a group of Grade 11 students on understanding soil science in Germany
orientation. This cycle of activities involved eliciting of pre-conceptions, restruc-
turing of student concepts during activities through comparison with original
ideas, clarification and exchange, construction and evaluation of new ideas, and
finally application, and review of new ideas. She uses five concrete steps in this
cycle (Drieling, personal communication):
1. Students imagine the ground under their feet and create a labeled sketch.
2. They next imagine that they shovel out part of the ground and note what they find.
26
Table 10 Initial understandings of soil by secondary students in three countries

Reference Country Students Probe used Topics studied Initial understandings
Drieling 2006, 2008 Germany Gr. 11 Guideline interviews and Structure and Soil is divided into layers of humus,
drawings components sand, or gravel, or soil is a place
of soil and its where animals live, or a habitat
functions for plans that also filters water.
Diagrams reveal students’ views
on soil erosion, acidification, and
hardening.
Happs 1981, 1984 New Zealand Gr. 12, University Interviews; (followed by a Makeup, origin, Students
and teachers’ Likert scale test survey depth, and age Knew the difference between soil
college of 16 items). of soils. Changes and dirt, in terms of some
students that soils components
undergo. Thought that soil come from a
multisource mechanism
Considered soils to be very, very
old
Thought the depth of soil was
between a few cms and 100 m
E- held a complicated view of the
soil-rock cycle
D. Hayhoe
Table 11 Treatment activities on soils for secondary students in three countries
Reference Country Students Probe used Treatment activities Results
Cattle et al. 1995 Australia High school, A computer program dealing with Feedback from universities and colleges
university the five main types of soil indicated that it has been used as a
students degradation was developed, teaching tool and as review material
(16–20 year) called “the Soil Stack.” in advanced courses
Drieling (2008) Germany Grade 11 (6th Interviews and A constructivist approach is used, The constructivist model is successfully
form) drawings where students first share their used to enable students to addess any
own pre-ideas, and they are misconceptions they have and be
exposed to hands-on examina- open to restructuring their ideas
tion of soil components and

soil profiles
Moebius-Clune US 48 Gr. 10 students 13 multiple 14 hands-on inquiry lessons on Pre-post gains went from 63% to 80%.
and Elsevier working in choice items water runoff and infiltration (Because students were from an
2008; small teams and 4 short into soils, with student teams agricultural area, many of them
Moebius- answer items creating their own questions scored very high on the pre-test.)
Clune et al. and doing their research
2011
27
28 D. Hayhoe
3. They then use a spade or an auger to expose a soil profile, performing appropriate
examinations on each horizon
4. They collect samples from the soil horizons and conduct lab tests such as pH.
5. They put together their results and draw conclusions
6. They compare and contrast their findings with the ideas they had initially.
The work of Moebius and Elsevier (2008) and Moebius-Clune et al. (2011)
involved 48 secondary school students, engaging in 14 hands-on inquiry lessons
concerning water runoff and infiltration into soils. Students asked their own
research questions, which included “questions assessing the influence of com-
paction, vegetation, rock content, particle size, slope, and prior water content
among others on runoff and infiltration partitioning.” Students completed a
series of worksheets to guide their inquiry, made journal entries, answers a
series of questions addressing the experiment and making connections to real-
world issues. (For the unit website, see Moebius and Elsevier 2008). The pre-
post test consisted of 13 multiple-choice questions from old multiple-choice
Regents exams (New York), and four short answer questions. Because the stu-
dents were from an agricultural area, many of them performed very well on the
pre-test.
In addition to answering pre-post tests, student teams also presented final proj-
ects related to their own inquiry research on runoff and infiltration. The distribu-
tion of the final project scores was bimodal. On the high end, a group of eight
students put in extra effort and moved beyond the majority. On the low end, a
number of projects had obviously been given little effort. Interestingly enough,
there was no correlation between test scores (13 multiple choice and 4 short
answer), and final project scores, although there was a negative correlation
between gains in test scores and final project scores, suggesting that the two meth-
ods of evaluation assessed different skills. Students also completed 11-item sur-
veys assessing their interest in the project. Scores were high on most items
(showing their enthusiasm and interest), and on two items they were very high.
Students overwhelmingly said they enjoyed this kind of research more than the
typical secondary school laboratory experiment, and that they learned to work in
a team as research scientists do.
The studies by Happs (1981, 1984) showed that secondary students undoubt-
edly approach the subject of soils with a more advanced initial understanding
than elementary students. The work of Drieling (2008) illustrates how the con-
structivist model of learning can be successfully combined with hands-on soil
profile investigations to enable senior secondary students to radically change
their ideas of soil. The work of Moebius-Cline et al. (2011) indicates that stu-
dents can indeed benefit from an intensive inquiry activities on particular soil
topics such as water runoff and infiltration. Other than these two studies, little
work has been done with secondary students in general. (See below for work
with students in specialized agricultural programs at the secondary school
level.)
2.4 Secondary School Teachers’ Knowledge of and Comfort

Level with Soils
There are no published studies I am aware of concerning initial understanding of

soil science by secondary school science teachers. Several studies have reported on
the readiness and comfort level of secondary school teachers (agriculture, social
studies, science) to teach about soils. Four of these are summarized here.
First, Puk and Behm (2003) studied the readiness of secondary school science teach-
ers to teach about environmental topics including soils. They sent out 500 surveys to
secondary school science and geography teachers across the province, and received
back 226 completed surveys. Results indicated that teachers often did not teach the
environmental components that had been infused into of the mainline science and
geography courses. Reasons included lack of time and lack of knowledge on the part of
the teacher. Soil concepts, in particular, were not taught by the majority of teachers.
Van Meter and Santucci (1990) were concerned about how frequently state soil
surveys (used in planting, land management, and by conservationists) were used by
secondary school agriculture and geography teachers across Indiana, an agriculture-
focused state. He sent out 184 surveys to all secondary schools, and received back
145 completed ones. Survey analysis indicated that while secondary school teachers
of agriculture were familiar with the soil surveys and most of them used the surveys
in class, only a minority of geography teachers knew about them and very few of
these made use of them in their classes. Interestingly, results were the same for
urban and rural areas. The researchers concluded that soil surveys were sill primar-
ily associated with agriculture and farm planning needs.
Wingenbach et al. (2007) studied the knowledge and comfort levels of preservice
agriculture teachers in Texas across the content topics of agricultural mechanics,
employability characteristics, agriculture and the environment, animal science, plant
and soil science, agricultural business management, soils and soil formations, and
food science. They found that these levels were considered adequate for all topics
except that of soils. Given what we know about the paucity of interest in soils and soil
education in elementary and secondary education, this might not be surprising,
although for the state of Texas soils do feature in the curriculum (Table 1).
Finally, Houck and Kitchel (2010) researched the content knowledge of preservice
agriculture teachers in Kentucky. In contrast to the previous findings of Wingenbach
et al. (2007), this study indicated that the teachers scored the highest on the topic of plant
and soil sciences, compared with animal sciences, agricultural engineering, agricultural
economics, and other agricultural social sciences. They note, however, that the students
had more course preparation in plant and soil sciences than in any of the other topics.
In summary, a handful of studies suggest that secondary school geography and
science teachers pay little attention to soil science, even though it may have rele-
vance to some of their curricular topics. With regard to preservice agriculture teach-
ers specifically, studies done in Kentucky and Texas give conflicting results as
to how well prepared they are to teach soil science at the secondary school level,
30 D. Hayhoe
compared with the other topics in their curriculum such as animal science, food sci-
ence, and agriculture management and engineering.
3 Resources Available for Elementary and Secondary

Soils Education
There are many excellent resources available in English for teaching soils at the
elementary level. These include complete equipment kits, extensive course manu-
als, articles from teacher journals, and websites with activities. At the middle school
and secondary school level, resources are primarily available in the form of journal
articles and websites.
3.1 Equipment Kits and Resource Manuals

for Teaching a Primary Soils Unit
3.1.1 Complete Equipment Kits
Many educational districts across North America have “science kit distribution
systems” that are able to provide primary teachers (K-4) with a complete soil sci-
ence kit for a month or more. When the teacher is finished with the kit, a staff of
technicians refurbish the kit and send it on to another teacher. Although complete
equipment unit kits cost between $500 and $1,000, the price per individual use is
relatively low. It contains all the equipment needed to lead a class through a series
of engaging hands-on activities with soil (sand, clay, silt, loam, magnifying glasses,
filters, seeds, trays, paper towels, etc. etc.). Most kits also come with a training CD
for the teacher, a manual of detailed class lessons, activity cards or “black line
masters” that can be used for each experiment, and assessment strategies and activ-
ities. The two kit programs described below (STC, FOSS) are ones that I have had
personal experience with and can attest to their quality.
STC Soils. The National Science Resources Center (NSRC) STC unit kits are avail-
able in English, Spanish, and Swedish for a cost of US $480. This kit is aimed at
the Gr. 2 level, although it has been used successfully in Gr. 3 classrooms across
Toronto. The unit includes 16 detailed hands-on lessons for students to work
through, using personal workbooks and comes with a teacher’s guide, a Teacher’s
Tools CD, 16 reusable student guides, and materials for a class of 32. (See http://
www.carolinacurriculum.com/STC/Elementary/Soils/index.asp, accessed on April
15, 2012). A sample lesson is available for download. The description on the NSRC
website is accurate: “Soils, a 16-lesson unit for second-graders, deepens children’s
awareness and appreciation of soil. Using simple tests, students learn to identify
sand, clay, and humus in soil. They also study how water affects different kinds of
soil. Through long-term experiments, they explore how roots and plants grow in
various soils and how, with the help of worms, old plants decompose and become
part of soil. Then, applying what they have learned, they investigate their own local
soil.”
FOSS Pebbles, Sand and Silt. The Full Option Science System (FOSS) is part of
the Lawrence Hall of Science. Like NSRC, FOSS kits cover all grades and strands
of Kindergarten to Grade 6 science. They are available for US $924 (http://www.
delta-education.com/fossgallery.aspx?menuID=2, accessed April 15, 2012). Like
STC Soils, this kit is aimed at Gr. 2 but is also suitable for Gr. 1–3. In addition to
the complete equipment the kit includes student videos and an up-to-date website
with pages for parents, teachers, and students (http://fossweb.com/modulesK-2/
PebblesSandandSilt/index.html, accessed on April 15, 2012). Unlike STC Soils,
the FOSS kit has students investigating rocks and minerals, leading them to see
where sand and clay come from, and then using them to construct and build objects,
before it concludes with soil explorations, where students put together and take
apart soils, are introduced to humus as an ingredient in soil, and compare home-
made and local soils.
3.1.2 Teacher Resource Manuals
Soils in the Environment (Andrews et al. rev. 2009). This 75-page, Gr. 3 resource
manual, with 25 laminated student activity sheets, includes 23 lessons on almost
every aspect of soil science appropriate for primary students. Topics include the
composition of soil, soil and water, characteristics of soil components, infiltration,
runoff, and sedimentation in water, soil profiles, organisms that live in the soil,
growth of plant roots in soil, and using composters. (Contact the author for avail-
ability at www.bill-andrew.com, accessed on April 15, 2012).
Dig in! Hands-on Soil Investigations (NSTA 2001). The National Resources
Conservation Services (NRCS) and the National Science Teachers Association
(NSTA) created this 129-page integrated resource, for elementary science teach-
ers and supervisors. It is aimed at the Kindergarten to Gr. 4 level. Each lesson
follows a five-step learning cycle – perception (30 min), exploration (30 min),
application (30 min), evaluation (15–30 min), and optional extensions (30 min
each) – and includes one or more student activity sheets (black line masters).
Topics addressed in the 12 lessons include components of soil (sand, clay, and
silt), formation of soil, soil layers, plants and animals that live in the soil, amount
of soil on the Earth and its use in food production, needs of plants growing in
soil, micro-organisms that live in the soil, food chains, worms, effects of water
and wind erosion on soil, soil scientists, creating a school garden. The book is
available in print (US $22) or electronic download (US $16) form NSTA (http://
www.nsta.org/recommends/ViewProduct.aspx?ProductID=12309, accessed on
April 15, 2012).
32 D. Hayhoe
3.1.3 Summary
Complete equipment kits are ideal for primary teachers who are able to focus a
unit of science teaching on soil. As they are expensive and require constant refur-
bishing, a central system is necessary to maintain science kits at the school, dis-
trict, or regional level. Of the two kits reviewed here, the STC Soils is more
focused on soils, and works very well for a curriculum unit only about soils.
FOSS Pebbles, Sand and Silt is better for an integrated rocks-soil curriculum unit
at the Gr. 1–3 level.
A complete teacher resource manual that includes a comprehensive series of les-
sons on soil science can also be a valuable resource. Of the two reviewed here,
Andrews’s Soil in the Environment contains more soil science. Although aimed at
Grade 3, it would also be suitable for the junior level (Gr. 4–6). NSTA’s Dig in!
Hands-on Soil Investigations is as extensive a resource, but is more appropriate for
the lower levels of primary school. It makes excellent use of the 5-E learning cycle
and requires less teacher background knowledge.
Other excellent books include Soils: Get the Inside Scoop (Soil Science Society
of America 2011), and Brown and Dickinson’s Earth Science: A Multifaceted
Investigation of Soil. (Zephyr, 1994).
3.2 Short Articles on Soil for Elementary, Middle School,

and Secondary School Teachers
Many excellent articles have been published in non peer-reviewed teacher journals
and other science activity journals with helpful ideas or perspectives on teaching
about soil science concepts at the elementary (Kindergarten to Grade 6), middle
school (Grades 7–9), or secondary (Grades 10–12) levels. These are generally
available for download from websites of libraries that subscribe to the journals
(Table 12). These articles should be considered supplementary to the use of unit
resources for teaching about soils, especially at the elementary level. For middle
school and secondary school teachers and soil educators (including extension), the
articles provide many good ideas.
3.3 Websites and Other Electronic Resources for Soil Education
Table 13 lists European and American websites on soil education. Many of the
European links are in the European Soil Data Centre inventory of educational material
(accessed at http://eusoils.jrc.ec.europa.eu/Awareness/inventory.cfm on April 1,
2012). Many of the American links are on the US Department of Agriculture website
for Soil Education (accessed at http://soils.usda.gov/education/ on April 15, 2012).
Table 12 Soil articles for elementary, middle school (Grade 7–9), and secondary (Grade 10–12) teachers
Grade level Title Description Journal Author(s) Date
Gr. 1–6 Boden (Soil) Special issue of a journal for Grundschulunterricht Various April 2007
primary teachers in (Translated:
Germany devoted to Soil Primary Education)
Gr. 1–2 “The radish party” The nature of soil, and its Science and Children Jeff Piotrowski, Tammy Oct. 2007, p 41–45
importance and relevance of Mildenstein, Kathy
soil organic matter for Dungan, Carol
young students Brewer
Gr. 1–2 “Second-grade soil Inviting a professional soil Science and Children Lori Gibb Nov/Dec 2000, p 24–28
scientists” scientist into a Gr. 2 class
working with the STC Soils
unit led to real inquiry
Gr. 1–6 “This land is your Hands-on activities with readily Science and Children Ann Kennedy, Tami Dec. 2006, p 22–26
land” available materials help L. Stubbs, Jeremy
students learn how to C. Hansen
prevent soil erosion
Gr. 1–6 “Is your soil sick?” Students explore the testing of Science and Children Donna R. Sterling April 2012, pp. 51–55
soil, learn about plant needs and Dori L.
and how to analyze results Hargrove
Gr. 1–8 “Synthetic soils: Students put together a mix of Science Activities Peter Veronesi Summer 1996, pp. 27–33
mixing the bad soil components and waste to
with the good” see how different plants grow
Middle “Science sampler: The Investigating the texture of soil, Science Scope Gary Bingham Oct 2010, pp. 63–68
school Science of soil and seeing how it has an
textures” effect on many different
kinds of activities
Middle “Hands-in science” Students study backyard soil to Science Scope Vickie Furlough, April 1997, pp. 16–17
school make a county soil map, and Amy Taylor, and
measure soil moisture and Scott B. Watson
porosity
33
(continued)
Table 12 (continued)
34
Grade level Title Description Journal Author(s) Date

Middle “Life in a teaspoon of Students use hand lenses or mic- Science Scope Shirley Foster Fields Feb. 1993, pp. 16–18
school soil” roscopes to find fungi in soil
Middle “Soil is more than just Students discover that soil is Science Scope Carrie Taylor and April 2010, pp. 70–74
school dirt” made of air, water, organic C. John Graves
material, and different
mineral grain sizes (sand,
silt, and clay), and has a
great diversity of organisms
living in it
High school “Soil searching: Students conducting chemical The Science Teacher Ann C. Kennedy, Feb. 1995, pp. 34–38
dishing the dirt on tests on various soils for Karen L. Smith,
microbes” their level of microbial Raymond L. Neff
activity
High school “Soil Testing: Dig in!” Students examine soil samples Journal of Chemical Linda N. Fanis and Feb. 2006, pp. 240A-B,
for physical appearance, Education Erica K. Jacobsen vol. 83, no. 2
water-holding capacity,
sedimentation, and pH
High school, “Quick, easy method This is a research article that Journal of Natural Thomas E. 2006, Vol. 25,
college to show living soil describes in detail useful Resources and Life Loynachan pp. 202–208
organisms …” ways to show the details of Sciences Education
microorganisms to high
school and college students
High school “Fungi: Strongmen of Students measure the weight of American Biology Patricia D. Morrell Jan. 1999, Vol. 61, No. 1,
the Underground” strings exposed to fungi in Teacher and Jeffrey pp. 54–55
wet soil, to develop a more J. Morrell
complete understanding of
fungi
High school “Measuring the Introduces a range of simple School Science Paul Perkins June 1994, Vol. 75,
physical properties ideas about soil physical Review No. 273, pp. 82–83.
D. Hayhoe
of soils” properties using a minimum

of apparatus
Table 13 Websites with soil education resources

URL Origin and content
European resources
http://www.al.fh-osnabrueck.de/fileadmin/ Media catalogue for the introduction of soil-related
users/30/upload/Bowi_Medienkatalog_ topics in school teaching in Germany
2009/medienkatalog_starten.html
http://www.bgr.bund.de/bodenunterricht Links to basic soil information and student
worksheets in the German language for
different levels and age groups
http://www.cienciadelsuelo.es/ Spanish resource on soil science, also available in
English. It is a multimedia- interactive program
with different modules that outline the study of
soil components and soil genesis
http://ec.europa.eu/environment/soil/pdf/ “Soil biodiversity: An excellent way to raise soil
Broll.pdf awareness,” a presentation in English, by Broll
(2010), on various aspects of soil education and
outreach in Europe
www.infovek.sk Educational resources for Slovakia
www.let-group.com Slovenian language website on the environment,
including soil, for primary, secondary, and
university students
http://www.macaulay.ac.uk/news/ The Dirt Doctors: Uses cartoon characters and
dirtdoctors/ humour to represent different soils. Comparing
human and soil health is an underlying theme.
Macauley Land Use Research Institute
www.soil-net.com Elementary and secondary educational resource,
supported by the British Society of Soil
Science. It uses cartoons at the primary level
(age 5–11) and informational text at the
secondary level (age 11–16).
American resources
http://archive.fieldmuseum.org/ Extensive soil resources for teachers and students
undergroundadventure/teachers/ for Kindergarten to Grade 8, by the Field
soil_biodiversity.shtml Museum in Chicago
www.dirtthemovie.com Tells the story of soil, “Earth’s most valuable and
underappreciated source of fertility–from its
miraculous beginning to its crippling degrada-
tion.” Available on itunes
www.doctordirt.org K-8 educational resources and activities on soils,
developed by Dr. Dirt, i.e., Clay Robinson, soil
scientist in West Texas
http://extension.usu.edu/aitc/lessons/index. Activities used in extension, by the Utah State
cfm University
http://nacdnet.org/education/resources/ Soil education resources sponsored by the National
soils/ Association of Conservation Districts, USA
http://soil.gsfc.nasa.gov/ NASA resources on soil science education
https://www.soils.org/lessons Elementary and secondary educational lessons,
developed by the Soil Science Society of
America
(continued)
36 D. Hayhoe
Table 13 (continued)
URL Origin and content
http://soils.usda.gov/education/ Soil education resources, and websites by the US
Department of Agriculture, for elementary,
secondary, and college levels
http://tlc.howstuffworks.com/family/ Science Projects for Kids: Soil – 5 hands-on soil
science-projects-for-kids-soil- experiments for elementary students, by The
experiments4.htm Learning Company
4 Conclusion
Studies conducted across a broad range of student ages, years, and countries con-
clude that students of all ages tend to begin the study of soils with very little
understanding about its composition, formation, and origin, although they appre-
ciate its necessity for life, especially plant life. With the use of an extended set of
effective hands-on activities lasting over several weeks, however, children as
young as 5–6 years are capable of achieving a lasting change in their mind-set
about soils, as demonstrated by interviews, diagrams, and objective tests held
months after the activities. Students above the age of 9 years can develop a deeper
understanding of the three-dimensional nature of soil, and begin to understand its
formation process and age. Secondary students are capable of going further, and
often learn a little about soils in their secondary ecosystem unit. Like students,
elementary teachers arrive at the study of soil with some pre-existing understand-
ing. In-depth classes in preservice science methods courses can significantly
increase their conceptual understanding. Little research has been done on soil
understanding of secondary teachers. It is only taught in specific agriculture
courses at that level.
Despite the importance of soil, and the ability of students, even those of young age,
to learn about it, the science curriculum of most countries does not allot a regular unit
to soils, although the topic is often touched upon in ecosystem units found in the biol-
ogy part of the curriculum. Canada and some parts of the U.S. are an exception to this.
In those regions, students study the topic in detail at the Grade 2 or 3 level, and many
excellent equipment kits, teacher resource manuals, and teacher articles are available.
Websites on soil education are ubiquitous throughout Europe and America.
Although the research on elementary and secondary soil education is limited,
results consistently indicate that student exposure to information and hands-on
experiments related to soil science can encourage long-term improvements in
student understanding of soil science. Given the resources available, one of the most
effective things that people concerned about soil education can do is to ensure that
the topic finds a permanent place in their country’s national curriculum (usually sci-
ence, but in some cases geography), and that classroom teachers are given the nec-
essary training, resources, and support by the countries soil science societies to
teach it in an effective manner.
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Sustainable Agriculture and Climate
Changes in Egypt
Hassan R. El-Ramady, Samia M. El-Marsafawy, and Lowell N. Lewis
Abstract Egypt is one of the most populous countries in Africa. Most of Egypt
82.2 million people live near the banks of the Nile River, in an area of about
40,000 km2, where the only arable land is found. The large areas of the Sahara
Desert are sparsely inhabited. About half of Egypt’s residents live in urban areas,
with most people spread across the densely populated centers of greater Cairo,
Alexandria and other major cities in the Nile Delta. Egypt’s fertile area totals about
3.3 million ha, about one-quarter of which is land reclaimed from the desert.
However, the reclaimed lands only add 7% to the total value of agricultural produc-
tion. Even though only 3% of the land is arable, it is extremely productive and can
be cropped two or even three times annually. Most land is cropped at least twice a
year, but agricultural productivity is limited by salinity, which afflicts an estimation
of 35% of cultivated land, and drainage issues. Climate change is a natural phenom-
enon, but humankind has drastically altered the process. Climate change has the
potential to affect agriculture through changes in temperature, rainfall timing and
quantity, CO2, and solar radiation. Agriculture can both mitigate or worsen global
warming. Some of the increase in CO2 in the atmosphere comes from the decompo-
sition of organic matter in the soil, and much of the methane emitted into the atmosphere
H.R. El-Ramady (*)

Soil and Water Sciences Dept, Faculty of Agriculture, Kafrelsheikh Uni,
33516 Kafr El-Sheikh, Egypt
e-mail: hassanelramady@rocketmail.com; ramady2000@gmail.com;
hassan.elramady@agr.kfs.edu.eg
S.M. El-Marsafawy
Soil, Water and Environment Research Institute (SEWRI),
Agricultural Research Center, Giza, Egypt
L.N. Lewis
Emeritus Professor, University of California, Rambla de Catalunya 47,
08007 Barcelona, Spain

42 H.R. El-Ramady et al.
is caused by the decomposition of organic matter in wet soils such as rice paddies.
Egypt’s agricultural development has been constrained by, among other factors, the
need to conserve scarce natural resources, the pressures of rapid urbanization, the
onslaught of the desert, and, not least important, technological limitations and
restrictive economic structures.
The major conclusions are (1) due to increasing recognition of climate change,
agriculture in Egypt is increasingly supporting issues of sustainable agricultural
production systems, and (2) most effects of climate change on sustainable agricul-
ture in Egypt could be changed through mitigation and adaptation.
Keywords Sustainability science • Climate changes • Sustainable agriculture

• Egypt
Abbreviations
BC Before Christ
CAPMAS Central Agency for Public Mobilization and Statistics
GDP Gross Domestic Production
GCOS Global Climate Observing System
GHGE Greenhouse Gases Emissions
IPCC Intergovernmental Panel on Climate Change
EEAA Egyptian Environmental Affairs Agency
INC Initial National Communication
MDGs Millennium Development Goals
MSL Mean Sea Level
NGOs Non-Governmental Organizations
SNC Second National Communication
SRU Strategic Research Unit
UNDP United Nations Development Programme
UNESCO United Nations Educational Scientific and Cultural Organization
UNFCCC United Nations Framework Convention on Climate Change
WMO World Meteorological Organization
1 Introduction
Egypt lies on the northeastern side of Africa, bordered on its northern coast by the
Mediterranean Sea and on its eastern coast by the Red Sea. It comprises an area of
about 1 million km2, made up as follows: Nile valley and delta about 4% of the total;
Eastern desert area about 22%; Western desert area about 68%; and the Sinai
Peninsula area about 6%. The share of Nile water in Egypt is 55.5 billion m3 year−1,
representing 76.7% of the country’s available water resources; desalinated seawater
Sustainable Agriculture and Climate Changes in Egypt 43
comprises only 0.08%. Total groundwater plus treated groundwater is 20.65 billion
m3 year−1 (28% of available water resources), but it cannot be added to Egypt’s share
of water as it is a reused source (CAPMAS 2009).
Awareness and concern for problems related to environmental quality are grow-
ing at a steady pace: climate change, biodiversity, soil fertility decay and above all
food quality and pollution are everyday subjects for debates and discussions. The
complexity of the problems and the uncertainty about many basic data quite often
make discussions inconclusive; even indications issued by scientific authorities are
sometimes misleading, and the problems are exacerbated by the frequent influence
of ideological positions (Wu and Sardo 2010).
Agriculture production has to increase by 70% within 2050 in order to keep pace
with population growth and changing diets. However, this production increase will
have to be achieved in a way that preserves the environment and reduces the vulner-
ability of agriculture to climate change. Agriculture will furthermore need to mini-
mize the emissions of greenhouse gases, pesticides and plant nutrients like nitrogen
and phosphorous to the environment (Aune 2012).
The main role of agriculture is to produce food for a growing population.
However, this production has to be achieved in an environmentally friendly way that
minimizes the external effects of agriculture related to the emission of green house
gases, the release of nitrogen and phosphorous to the environment and the use and
accumulation of harmful pesticides in nature. Agriculture will also need to adapt to
climate change including more extreme weather events. In principle, there are three
pathways for agricultural development: conventional agriculture, organic agricul-
ture and conservation agriculture. These pathways have different approaches for
addressing the above issues (Aune 2012).
Despite its prediction 100 years ago by scientists studying CO2, manmade cli-
mate change has been officially recognized only in 2007 by the Nobel Prize
Committee. Climate changes since the industrial revolution have already deeply
impacted ecosystems. The lesson from the climate change story is that humans do
not learn from scientists until it really hurts. Furthermore, all society issues cannot
be solved anymore using the old, painkiller approach because all issues are now
huge, linked, global and fast-developing. In that respect, actual society structures
are probably outdated. Here, agronomists are the most advanced scientists to solve
society issues because they master the study of complex systems, from the molecule
to the global scale. Now, more than ever, agriculture is a central point to which all
society issues are bound; indeed, humans eat food (Lichtfouse 2009).
More than 100 years ago, the Nobel Prize winner Svante Arrhenius (1859–1927)
estimated that a doubling of atmospheric CO2 concentration would cause a tempera-
ture rise of about +5−6°C (Arrhenius 1896). Remarkably, his crude estimate is
higher but not largely different from the +2.0−4.5°C rise now estimated by the
Intergovernmental Panel on Climate Change (IPCC 2007). Combining his calcula-
tions with existing work suggesting that the burning of fossil fuels could significantly
alter the concentration of carbon dioxide in the atmosphere (Hoegbom 1894),
Arrhenius later became the first person to predict the possibility of man-made global
warming (Arrhenius 1908; Weart 2008; Lichtfouse 2009).
2 Sustainable Agriculture in Egypt
Some researchers define sustainable agriculture primarily as a technical process.

Altieri (1989) defined sustainable agriculture as a system, which should aim to
maintain production in the long run without degrading the resources base, by using
low-input technologies that improve soil fertility, by maximizing recycling, enhanc-
ing biological pest control, diversifying production, and so on. The technological
and to a lesser extent economic dimensions of sustainable agriculture have tended
to be privileged while the social dimension has been neglected. As a result sustain-
able agricultural has suffered from limited adoption (Karami and Keshavarz 2010).
A sustainable farming system is recognized as a system that maintains the
resource base upon which it depends, relies on minimum of synthetic inputs, man-
ages pests and diseases through internal regulating processes, and can recover from
the human disturbance caused by agricultural practices, i.e., cultivation and harvest
(Altieri 1995). Sustainable agriculture is farming systems that are maintaining their
productivity and benefit to society indefinitely (Lichtfouse et al. 2009).
Despite the diversity in conceptualizing sustainable agriculture, there is a consen-
sus on three basic features of sustainable agriculture: (1) maintenance of environ-
mental quality, (2) stable plant and animal productivity, and (3) social acceptability.
Consistent with this, Yunlong and Smith (1994) have also suggested that agricultural
sustainability should be assessed from ecological soundness, social acceptability,
and economic viability perspectives. “Ecological soundness” refers to the preserva-
tion and improvement of the natural environment, “economic viability” to mainte-
nance of yields and productivity of crops and livestock, and “social acceptability” to
self-reliance, equality, and improved quality of life (Karami and Keshavarz 2010).
In conclusion, sustainable agriculture is defined simply as farming systems that
are maintaining their productivity and benefit to society indefinitely. This sustain-
able agriculture has three basic features, e. g. maintenance of environmental quality,
stable plant and animal productivity, and social acceptability.
2.1 Definition of Agriculture
What is agriculture? It is the first point to clarify and there is of course general
agreement about the sorts of things, people, plants, and animals that can be called
agricultural, but this is not good enough if we are seriously interested in topics such
as the role of science in agriculture, the role and importance of agriculture in the
world, and how agricultural efficiency can be improved (Speeding 1988). Not many
attempts have been made to be more precise and it is quite difficult to arrive at a
definition that is both useful and specific. One of the useful definitions is phrased by
Speeding (1988, 1996) as follows: “agriculture is an activity of Man, carried out
primarily to produce food, fiber and fuel, as well as many other materials by the
deliberate and controlled use of mainly terrestrial plants and animals” (Karami and
Keshavarz 2010).
Therefore, agriculture also called farming or husbandry is the cultivation of ani-

mals, plants, fungi, and other life forms for food, fiber, biofuel and other products
used to sustain life. Agriculture was the key development in the rise of sedentary
human civilization, whereby farming of domesticated species created food surpluses
that nurtured the development of civilization. The study of agriculture is known as
agricultural science.
2.2 Sustainability Science
Sustainability is the core element of government policies, university research proj-

ects, and extension organizations worldwide. Yet, the results of several decades of
attempt to achieve sustainable agriculture have not been satisfactory. Despite some
improvement conventional agriculture is still the dominant paradigm. Sustainability,
climate change, and replacing fossil fuels with renewable energy are relatively new
challenges for agriculture. Sustainability is not only a challenge in itself, but also a
new worldview, a paradigm, which has changed our understanding of agriculture.
Sustainable agriculture as a concept has emerged to address the challenges that are
facing modern agriculture. Across all literatures, two broad paradigms of sustain-
ability are identifiable: one supporting a systems-level reconstruction of agricultural
practice to enhance biological activity, and the other adopting a technological fix, in
which new technologies inserted into existing systems can improve sustainability
outcomes (Karami and Keshavarz 2010).
It could be selected from some of the major ideas of sustainability science that
contributed to the development of sustainability science from a much larger set,
beginning first with Alexander von Humboldt’s dream of understanding the unity of
nature. This was followed by George Perkins Marsh’s vision of nature as modified
by human action. Then much later, the International Union for the Conservation of
Nature (IUCN) linked nature and human development, which led to the World
Commission on Environment and Development, and culminated in the US National
Academy of Science (NAS) report of Our Common Journey and the call for a sus-
tainability science (Kates 2012).
Sustainability science could be defined as follows “an emerging field of (transdis-
ciplinary) research dealing with the interactions between natural and social systems
… how those interactions affect the challenge of sustainability: meeting the needs of
present and future generations while substantially reducing poverty and conserving
the planet’s life support systems” (Weinstein and Turner 2012). Balancing human
needs with the ability of ecosystems to provide the goods and services that we all
depend on is a fundamental formula for the global sustainability transition (Fig. 1).
Equilibrium can be attained either by increasing these goods and services or by
reducing our consumption of them, or in today’s world, both! Any solution to the
emerging conflicts arising on the path to long-term sustainability will, in part,
require the integration of the biophysical and social sciences into a new transdisci-
plinary science that we refer to as “sustainability science”.
The Sustainability Equation
Ecosystem
Goods &
Services Human
Input Output
Ecosystem Progress &
Integrity Quality of
Sustainable Life
Ecocentrism Development Anthropocentrism
o Biodiversity o Hazard Mitigation
o Ecosystem Resilience o Buffering Core Tech.
o Habitat Complexity o Ecosystem Health
o Few Invasive Taxa Preservation, conservation, o Environmental Justice
Restoration, Ecosystem-Based o Recreation & Aesthetics
Management, Adaptive o Engineering Resilience
Management
Human Progress & Quality of Life ≤ Ecosystem Goods & Services
Fig. 1 The “Sustainability Equation” balancing human needs with ecosystem integrity (Adapted
from Weinstein and Turner 2012). Note the increase of the inputs (ecosystem integrity) through
ecosystem goods and service follows by increasing the outputs (human progress and quality of life
and finally sustain the global development)
Therefore, sustainability science has emerged in the twenty-first century as a new

academic discipline. This new field of science was officially introduced with a
‘Birth Statement’ at the World Congress ‘Challenges of a Changing Earth 2001’ in
Amsterdam. The name of this scientific field reflects a desire to give the generalities
and broad-based approach of “sustainability” a stronger analytic and scientific
underpinning. Sustainability science, like sustainability itself, derives some impetus
from the concepts of sustainable development and environmental science.
Sustainability science provides a critical framework for sustainability while sustain-
ability measurement provides the evidence-based quantitative data needed to guide
sustainability governance.
2.3 Sustainable Development
Sustainable development was defined as the development that meets the needs of
the present without compromising the ability of future generations to meet their
own needs. If needs are to be met on a sustainable basis, the Earth’s natural resources
must be well managed and enhanced. Also, it is defined as a development strategy
that manages all assets, natural resources and human resources as well as financial
and physical assets, for increasing long-term wealth and well-being. Development
is a value-laden word implying change that is desirable. It may be considered a vector
of desirable society objectives, the elements of which might include (Pearce et al.
1990): increase in real income per capita, access to resources, and a “fairer” distri-
bution of income. Sustainable development has been described as a path toward the
twin goals of social justice and environmental protection. It rejects policies and
practices that support current living standards by depleting the productive base,
including natural resources and that leaves future generations with poorer prospects
and greater risks than our own. In its broadest sense, the strategy for sustainable
development aims to promote harmony among human beings and between humanity
and nature. Bases for sustainable development include: (1) reliable scientific infor-
mation, (2) consensus on ethical principles, (3) hope for the future, and (4) consid-
eration of personal interest and incentives. “It could be managed as our resources
only if we know what we have and what we are doing to them. We need to agree on
the reasons for preserving and distributing resources. We also need assurance that
progress is possible and that we or our descendents will benefit from that progress”
(Cunningham and Saigo 1992) Also, Kassas (2004) stated that sustainable develop-
ment could be realized through three main bases: (1) social equality, (2) economic
efficiency and (3) environmental conservation. He added that it is the responsibility
of the governmental institutions, in collaboration with the international and regional
organizations, to implement programs aiming at the conservation of the natural
resources, e.g. the biodiversity, and to protect them against deterioration (Zahran
and Willis 2009).
In conclusion, sustainable development (SD) is a pattern of economic growth in
which resource use aims to meet human needs while preserving the environment so
that these needs can be met not only in the present, but also for generations to come.
The term “sustainable development” was used by the Brundtland Commission
which coined what has become the most often-quoted definition of sustainable
development as development that “meets the needs of the present without compro-
mising the ability of future generations to meet their own needs.” Alternatively,
sustainability educator Michael Needham referred to sustainable development “as
the ability to meet the needs of the present while contributing to the future genera-
tions’ needs” (Needham 2011). There is an additional focus on the present genera-
tions “responsibility to improve the future generations” life by restoring the previous
ecosystem damage and resisting to contribute to further ecosystem damage.
2.4 Sustainable Agriculture in Ancient Egypt
Ancient Egypt was an ancient civilization of Northeastern Africa, concentrated

along the lower reaches of the Nile River in what is now the modern country of
Egypt. Egyptian civilization coalesced around 3150 BC (according to conventional
Egyptian chronology) with the political unification of Upper and Lower Egypt
under the first pharaoh. The search for an ancient society that approached a sustain-
able balance with the environment must inevitably lead the environmental historian
to Egypt. The Egyptians were in charge of their own government and able to set
their own environmental policies from before 3000 BC to after 1000 BC. No other
ancient civilization lasted so long, while maintaining a stable pattern in its economy,
government, religion, and ecological viewpoints and techniques. Many historians of
Egypt remark upon the stability of Egyptian culture in a pejorative tone, attributing
a lack of change to traditionalism and absence of creative thought, as if stability
were only stagnation. But it will be suggested here that the stability of Egyptian
civilization was the result of the sustainability of Egypt’s ecological relationships
(Hughes 1992).
The many achievements of the ancient Egyptians include the quarrying, survey-
ing and construction techniques that facilitated the building of monumental pyra-
mids, temples, and obelisks; a system of mathematics, a practical and effective
system of medicine, irrigation systems and agricultural production techniques, the
first known ships, Egyptian faience and glass technology, new forms of literature,
and the earliest known peace treaty with Hittites. The sustainability of Egyptian
agriculture was made possible first of all by the annual flood of the Nile and the
deposition of fertile alluvial soil containing phosphorus and other minerals and
traces of organic debris brought down from the mountains and swamps of lands
further south. The Greek historian Herodotus, observing that the very soil of Egypt
had been formed by river sediment, pronounced Egypt the “gift of the Nile”. The
Egyptians were aware of this, for as an early inscription witnesses, the Nile supplies
all the people with nourishment and food. Second, they had a stable climate without
freezing or storms, and although there was little rain, the river supplied the water
needed. Their environment encouraged them to think of processes of nature as oper-
ating in predictable cycles. The Nile flooded its banks at the same time every year,
bringing moisture and new soil to the fields, and then subsided. The only fertile land
was what the river watered, both in the long, narrow cultivated valley floor of Upper
Egypt and in the broad, flat, fruitful Delta of Lower Egypt (Fig. 2; Hughes 1992).
Egypt left a lasting legacy. Its art and architecture were widely copied, and its
antiquities carried off to far corners of the world. Its monumental ruins have inspired
the imaginations of travellers and writers for centuries. A new-found respect for
antiquities and excavations in the early modern period led to the scientific investiga-
tion of Egyptian civilization and a greater appreciation of its cultural legacy. In spite
of Egypt’s remarkable success in maintaining sustainable agriculture, some envi-
ronmental problems appeared. One, ironically, was a result of the success of the
Egyptians in producing the ancient world’s most reliable food supply. The most
dependable system will fail with over-population. When population increased to
near a level that could be supported in a year of good harvest, an abnormally low
harvest would bring famine. Egypt suffered because fat years alternated with lean
ones, and population had its peaks and valleys as a result. Governmental officials
tried to even out fluctuations of supply and demand by storing surplus in good years
and distributing it when the harvest failed. The story of Joseph’s interpretation of
Sheep, goats, cattle, pigs and geese were raised Wall painting from 1200 BC showing an Ancient
from earliest times and supplied milk, wool, Egyptian ploughing his field
meat, eggs, leather, skins, horn and fat
A tomb relief depicts workers plowing the fields, Sennedjem plows his fields with a pair of oxen,
harvesting the crops, and threshing the grain used as beasts of burden and a source of food.
under the direction of an overseer
Fig. 2 Some agricultural practices photos from Ancient Egypt from different websites (www.
ancientegyptmagazine.com, www.wekipedia.com and www.experience-ancient-egypt.com/images/
ancient-egyptian-farming.jpg/ 28.6.2012)
Pharaoh’s dream, and his advice to build granaries to prepare for hard times, is a
reflection of the actual situation in Egypt. The store chambers of the Ramesseum,
built at the order of Ramses II, could easily have held 590,000 cu. ft. of grain,
enough to support 3,400 families for a year. In difficult periods, prices fluctuated
widely. In the 55 years between the reigns of Ramses III and Ramses VII, for exam-
ple, emmer wheat rose from 8 to 24 times base price. At times, famine relief had to
be distributed over wide territories. Even so, Egypt remained the breadbasket of the
ancient world, exporting wheat and barley with few interruptions (Hughes 1992).
Therefore, a combination of favorable geographical features contributed to the
success of ancient Egyptian culture, the most important of which was the rich fertile
soil resulting from annual inundations of the Nile River. The ancient Egyptians
were thus able to produce an abundance of food, allowing the population to devote
more time and resources to cultural, technological, and artistic pursuits. Land manage-
ment was crucial in ancient Egypt because taxes were assessed based on the amount
of land a person owned. Farming in Egypt was dependent on the cycle of the Nile
River. The Egyptians recognized three seasons: Akhet (flooding), Peret (planting),
and Shemu (harvesting). The flooding season lasted from June to September,
depositing on the river’s banks a layer of mineral-rich silt ideal for growing crops.
After the floodwaters had receded, the growing season lasted from October to
February.
2.5 Sustainable Agriculture in Modern Egypt
The total area of Egypt is about 1 million km2, most of which is under arid and
hyper-arid climatic conditions, and of which a small portion representing only 3%
is agriculturally productive. The six main agro-ecological zones in Egypt are the
Nile Valley including the fertile alluvial land of Middle and Upper Egypt, where the
main source of irrigation water is the Nile River. Agriculture production of Egypt is
mainly concentrated in this zone in addition to the Delta as follows:
1. The Nile Delta region, where the main source of irrigation water is the Nile River
as well. Together with the Nile Valley, the agriculture production in this zone
consists of about 6.6 million acres. Most of the soil in both areas is recent Nile
alluvium.
2. The reclaimed desert areas in the fringes of the Nile Valley, where the only source
of irrigation is the ground water.
3. North Coastal zone: including the coastal area starting from North-Western coast
moving eastwards to North coastal area of Sinai Peninsula, where there are no
reliable figures are available neither on ground water quantity or usage.
4. The Inland Sinai and the Eastern Desert, where the main source for irrigation is
the ground water.
5. The Western Desert including oases and southern remote areas, where the ground-
water is mainly extracted from the Nubian Sandstone and carbonate aquifers.
Although the Nile River streams through the Egyptian land, water is regarded as
a scarce natural resource, due to the rapidly growing population in Egypt, the lat-
ter’s limited quota of the Nile Water and the wide desert lands where the main drink-
ing and irrigation water resource is the underground water. Furthermore, the
Egyptian land suffers from different variations of degradation around the country,
depending on the region and the inhabitants (Afifi 2009).
According to report of Handoussa (2010) about situation analysis: key develop-
ment challenges facing Egypt, it could be concluded the following topics about the
sustainable agriculture in modern Egypt:
1. Issues of Sustainable Agricultural Development:
The link between environmental and sustainable agricultural and rural development,
enhancing food security, and reducing poverty is a central issue for achieving eco-
nomic and social development in Egypt. The contribution of the agriculture sector
in Egypt exceeds 13% of GDP and over 30% of employment opportunities.
Meanwhile, about 57% of the total population in Egypt lives in rural areas, where
poverty prevails. As such, enhancing sustainable agricultural and rural development
as a means to reduce poverty and food insecurity within the expected climate
changes is a prerequisite for sustainable social and economic development and

hence should be considered as a social and political priority for Egypt. In Egypt,
agriculture is recognized as a way of life and crucial for socio-economic develop-
ment, but if received the due attention, also as an engine for growth.
2. Rural Poverty and Food Security:
There is a strong correlation between economic growth and the reduction of hunger
and poverty as well as a strong link between poverty and food insecurity. Most of
the poor are either under-nourished or food insecure. Lower income households
spend a large share of their income to purchase food. They are particularly vulner-
able to variations in food prices and food scarcity. Nearly 70% of the poor or food-
insecure live in rural areas and a large share of these people depend very much on
agriculture for their food supplies (produced locally) and for generating incomes.
Economic diversification starts at the farm household, and agricultural and non-
agricultural development reinforce each other. Pro-poor policies and strategies must
emphasize food security, access to land and water, agriculture and rural develop-
ment. Large numbers of rural households depend on agriculture and farming (pro-
duction) but it is rarely the main income contributor. Farm incomes account on
average for about 25–40% of total rural income, agricultural related off farm
incomes account for an additional 20–35%, and non-farm revenues and wages
account for about 40% of rural household incomes.
3. Improving Irrigation Efficiency:
Pollution of waterways and groundwater due to domestic and industrial wastewater
and solid waste disposal is another problem that reduces the availability of appropri-
ate quality water for use. The competition between development sectors on water
and between users of the same sector is growing and is expected to create water
conflicts. The number of irrigation water complaints due to water shortages is known
to have increased recently. Due to the limited water resources, the irrigation water
shortages are exacerbated by illegal water intakes and violations in cultivating more
than the allowable areas of high water consumptive crops, such as rice. Treated
wastewater is expected to be the renewable water resource for agriculture expansion
in the future, if no additional share of Nile waters is mobilized, and the existing
Egyptian code for treated wastewater reuse in agriculture will need to be properly
implemented.
4. Depletable Energy Sources:
Hydrocarbons such as oil and gas represent over 90% of Egypt’s energy resources,
a situation that is expected to continue for at least the next 20 years, with natural gas
slowly replacing crude oil. Egypt’s reserves have been increasing at an average of
5% per year during the last 5 years and natural gas reserves accounted for most of
this growth, while crude oil and condensate reserves remained relatively constant.
Domestic consumption however, has been rapidly increasing as a result of ongoing
economic growth. Recent and forecasted consumption patterns signal an alarm to
those responsible for the country’s future economic development and energy security.
Energy efficiency and renewable energy resources are expected to play a critical
role in facing this challenge.
5. Climate Changes:
The energy sector is the main source of greenhouse gas emissions with 92% of the
country’s energy demand met by using fossil fuels. It is worth noting that Egypt’s
greenhouse gas emissions are relatively limited (0.7% of global greenhouse gas),
but they grew to 193 million tons of CO2 equivalent in 2000 from 116 million tons
of CO2 equivalent in 1990 (Egypt’s initial national communication/second national
communication). However, Egypt is subject to potential impacts of climate
change, including sea level rise, inundation of the low lying lands in the Nile
Delta that could reach 10–12% of the total area, impacts on water resources and
agricultural productivity and associated social and economic effects. Moreover,
57% of the Egyptian population lives in rural areas, considered more vulnerable
to climate change, with an expected shortage of basic food items. The increased
concentration of greenhouse gases in the atmosphere are causing disruptions in
climate systems and the broad impacts can be divided into two main groups
(Table 1).
6. Impact of Climate Change on Health:
Climate change is also closely linked to the health sector, with expected
increases in morbidity and deaths due to non communicable diseases, disasters,
and vector-borne diseases. There are some efforts to reduce the impacts of cli-
mate change in some sectors. Currently, however, there are no efforts to cope
with the different direct and indirect impacts of climate change on health.
Moreover, building institutional capacity must be pursued vigorously in order
to deal adequately with the necessary adaptation measures, providing the
country with both a strategy and a trained capacity to implement the required
measures.
7. Egypt 2050 – The Need for a National Urban Development Plan:
If Egypt’s continued high rate of natural population increase is not reduced it could
result in a population of 140 million inhabitants by 2050. This challenge calls for
the need to develop a new vision for Egypt that mainly aims to:
• Achieve balanced urban development by focusing on the development of small
and medium-sized cities, especially in Upper Egypt, to eradicate poverty and
improve the socio-economic status within those deprived areas;
• Redefine the roles of existing poles such as Cairo, Alexandria, Port Said and others
in an overall policy framework that focuses on building on the regional competi-
tiveness potentials of each pole and how it can be integrated within the overall
new urban development policy;
• Increase the inhabited area from 5.5% to 15% of the total area during the next four
decades through establishing new urban centers that are well connected with
efficient road networks and national public transportation systems. Each center
would work as a catalyst for the development of its surroundings and attracting a
defined number of the increasing population with a clear economic basis and
activities;
Table 1 Egypt at a glance from 1988 to 2008 through some indicators: the key economic ratios
and long-term trends, structure of economy (average annual growth and gross domestic production),
trade and balance of payment (Adapted from UNDP 2011)
Item 1988 1998 2007 2008
Key Economic Ratios and Long-Term Trends
GDP (US$ billions) 35.0 84.8 130.5 162.3
GDP (average annul growth) 4.1 4.6 7.1 7.2
GDP per capita 2.0 2.7 5.1 5.2
Gross capital formation/GDP 34.9 21.5 20.9 22.5
Exports of goods and services/GDP 17.3 16.2 30.3 33.2
Current account balance/GDP −1.6 −2.9 1.7 0.5
Structure of Economy (% of GDP)
Agriculture 19.0 17.1 14.1 13.2
Industry 28.8 30.9 36.3 37.5
Services 52.2 52.0 49.6 49.2
Imports of goods and services 35.2 25.7 34.8 38.8
Structure of Economy (average annual growth)
Agriculture 2.9 3.4 3.7 3.3
Industry 6.0 4.9 7.9 10.3
Services 3.0 5.1 7.4 8.6
Imports of goods and services 2.8 11.4 28.8 26.3
Trade (US$ millions)
Total exports (free on board, fob) 3,274 5,128 22,018 29,356
Cotton 480 1,728 110 194
Other agriculture 354 103 10,223 14,628
Manufactures 961 1,885 7,519 10,932
Total imports (cost, insurance, freight, cif) 8,858 16,899 38,308 52,771
Food 1,254 3,193 2,671 3,927
Fuel and energy 2,148 2,188 4,336 10,001
Capital goods 2,188 4,801 9,845 11,871
Balance of Payments (US$ millions)
Exports of goods and services 7,225 13,502 39,428 53,277
Imports of goods and services 11,689 21,795 45,398 63,086
Resource balance −4,465 −8,292 −5,969 −9,809
Net income −161 1,213 1,177 1,360
Current account balance −545 −2,479 2,269 888
GDP gross domestic production, US$ United States dollar
• Deal with the potential link between climate change (including SLR, water scar-
city and desertification) and human mobility (including displacement) in Egypt.
Given its potential magnitude, environmentally induced migration could impact
adversely upon various human development issues (for example, rapid urbanization
and associated environmental health issues) and undermine the important progress
outlined in this report;
• Support the decentralization of management, planning and implementation.
In conclusion, agriculture production of Egypt is mainly concentrated in the Nile

Valley zone in addition to the Delta. About the sustainable agriculture in modern
Egypt, it could be concluded that the key sustainable development challenges facing
Egypt include the following topics: climate change, issues of sustainable agricultural
development, impact of climate change on health, rural poverty and food security, the
need for a national urban development plan, depletable energy sources and improv-
ing irrigation efficiency.
3 The Climate of Egypt
The land of Egypt occupies the northeastern part of the African continent. It is
roughly quadrangular, extending about 1,073 km from north to south and about
1,229 km from east to west. Thus, the total area of Egypt is a little more than one
million square kilometers (1,019 600 km2) occupying nearly 3% of the total area of
Africa (Abu Al-Izz 1971). Egypt is bordered on the north by the Mediterranean Sea,
on the south by the Republic of Sudan, on the west by the Republic of Libya and on
the east by the Gulf of Aqaba and the Red Sea.
Egypt extends over about 10° of latitude, being bounded by Lat. 22 °N and 32 °N,
i.e. it lies mostly within the temperate zone, less than a quarter being south of the
Tropic of Cancer. The whole country forms part of the great desert belt that stretches
from the Atlantic across the whole of North Africa through Arabia. Egypt is charac-
terized by a hot and almost rainless climate. The average annual rainfall over the
whole country is only about 10 mm. Even along the narrow northern strip of the
Mediterranean coastal land where most of the rain occurs, the average annual rain-
fall is usually less than 200 mm and the amount decreases very rapidly inland
(southwards). The scanty rainfall accounts for the fact that the greater part of Egypt
is barren and desolate desert. Only through the River Nile is a regular and volumi-
nous supply of water secured, coming from the highlands hundreds of kilometres to
the south. This is channeled by artificial canals over the narrow strip of alluvial land
on both sides of the river, the Fayoum Depression and the delta expanse. These
tracts of fertile land, covering less than 3% of the total area of Egypt, support a
dense population (Zahran and Willis 2009).
According to Said (1962), the average density of population in the agricultural
lands of Egypt is more than 600 persons km2, whereas in the vast desert areas, which
represent more than 97% of the total area, there is only one inhabitant/7 km2. The
River Nile, therefore, is a salient geographical feature that has shaped not only the
physical tracts of Egypt but also its history and the nature of its human settlements.
Herodotus (484–425 BC) states that “Egypt is the Gift of the Nile”. This is very true
as the Nile gave Egypt, out of all regions of the great North African Sahara, a fertil-
ity that made possible not only the development of the famed ancient agricultural
civilization, but also the growth of this civilization in peace and stability (Zahran
and Willis 2009).
3.1 Climate and Climate Changes
Climate refers to the characteristic conditions of the earth’s lower surface atmo-
sphere at a specific location; weather refers to the day-to-day fluctuations in these
conditions at the same location. The variables that are commonly used by meteo-
rologists to measure daily weather phenomena are air temperature, precipitation
e.g., rain, sleet, snow and hail, atmospheric pressure and humidity, wind, and sun-
shine and cloud cover. There is no internationally agreed definition of the term
“climate change”. Climate changes can refer to: (1) long-term changes in average
weather conditions (World Meteorological Organization usage); (2) all changes in
the climate system, including the drivers of change, the changes themselves and
their effects (Global Climate Observing System usage); or (3) only human-induced
changes in the climate system (United Nations Framework Convention on Climate
Change usage). Climate change as referred to in the observational record of climate
occurs because of internal changes within the climate system or in the interaction
among its components, or because of changes in external forcing, either for natural
reasons or because of human activities. It is generally not possible to make clear
attributions between these causes. Projections of future climate change reported by
IPCC generally consider the influence on climate of only anthropogenic increases
in greenhouse gases and other human related factors (IPCC usage) (FAO 2008).
According to the IPCC (2008), climate change is any “change in climate over time
whether due to natural variability or as a result of human activity”. It is general
consensus among IPCC researchers that increases in atmospheric concentrations of
greenhouse gasses (mainly CO2, CH4, N2O and O3) since pre-industrial times have
led to a warming of the surface of the earth. During the last 250 years, the atmo-
spheric concentrations of CO2, CH4 and N2O have increased by 30%, 145% and 15%,
respectively. The emissions are mainly due to the use of fossil fuels, but changes of
land use as well as agriculture are also major sources of emissions (Raberg 2008).
Therefore, climate change is a significant and lasting change in the statistical
distribution of weather patterns over periods ranging from decades to millions of
years. It may be a change in average weather conditions, or in the distribution of
weather around the average conditions i.e., more or fewer extreme weather events.
Climate change is caused by factors that include oceanic processes such as oceanic
circulation, variations in solar radiation received by Earth, plate tectonics and vol-
canic eruptions, and human-induced alterations of the natural world; these latter
effects are currently causing global warming, and ‘climate change’ is often used to
describe human-specific impacts.
3.2 Why the Interest in Global Climate Change?
The world population is projected to increase from about seven billion in 2011 to
9.2 billion in 2050. The current rate of increase is about six million per month, with
almost all growth occurring in developing countries where natural resources are
already under great stress. The Green Revolution technology led to the doubling of
food production between 1950 and 2010, with only a 10% increase in the area under
production (FAO 2010). However, meeting the food demand of the growing popula-
tion, rising standards of living, and changes in diet preferences will necessitate an
additional 70% increase in production between 2010 and 2050 (Burney et al. 2010).
Grain yields of wheat (Semenov 2009) and rice (Wassmann et al. 2009) are sensitive
to high temperatures (Lal and Stewart 2012).
Climate changes caused by the progressive anthropogenic emissions of green-
house gases is already affecting natural and human systems and sectors throughout
the world and the changes to date may be only inklings of profound changes to
come. Some contend that action on climate change should be delayed because of the
uncertainties surrounding the exact nature, extent, and rate of the portending
changes. Others believe that responding to climate change is now necessary pre-
cisely because of the uncertainties. In any case, the prospect of significant changes
in agroecosystems requires us to anticipate the potential impacts of climate change,
to study how farming regions and systems can adjust to those that are unavoidable,
and to determine how they can mitigate climate change so as to reduce its ultimate
effects (Hillel and Rosenzweig 2011).
From previous, it could be concluded that climate changes in different regions of
the world showed that it is likely to vary a great deal from place to place. For instance,
in some regions precipitation will increase, in other regions it will decrease. Not only
is there a large amount of variability in the character of the likely change, there is also
variability in the sensitivity of different systems to climate change. Different ecosys-
tems, for instance, will respond very differently to changes in temperature or precipi-
tation. There will be a few impacts of the likely climate change that will be positive
so far as humans are concerned. For instance, in parts of Siberia, Scandinavia or
northern Canada increased temperature will tend to lengthen the growing season
with the possibility in these regions of growing a greater variety of crops. Also, in
winter there will be lower mortality and heating requirements. Further, in some
places, increased carbon dioxide will aid the growth of some types of plants leading
to increased crop yields. However, because, over centuries, human communities have
adapted their lives and activities to the present climate, most changes in climate will
tend to produce an adverse impact. If the changes occur rapidly, quick and possibly
costly adaptation to a new climate will be required by the affected community. An
alternative might be for the affected community to migrate to a region where less
adaptation would be needed – a solution that has become increasingly difficult or, in
some cases, impossible in the modern crowded world.
3.3 Effects of Climate Change on Soil and Water Resources
Soil water balance has response to climate change and evaluation of soil water
change is one of the most important items of climate change impact assessment.
It is well documented also that a changing climate will affect soil and water resources
Channel Stability
Flooding
Stream Flow
Water
Water Supply Pollution
RUNOFF
Precipitation (Leaching)
Soil Moisture
Biomass Management
CO2 EROSION
Temperature
Soil Degradation Water Pollution
Solar Radiation
Indicates the pathways and effects investigated in this report
Fig. 3 Effects of climate change on soil and water resources (Adapted from SWCS 2003). Erosion
and runoff are among the most important factors influencing agriculture’s effects on soil and water
resources, precipitation would be a dominant factor affecting conservation outcomes and that soil
erosion and runoff would be particularly sensitive to changes in precipitation
on agricultural land in many ways. Will the effect of climate change on soil and
water resources on agricultural land be large enough to warrant changes in world
conservation policy or practice? Climate change will affect soil and water conserva-
tion through multiple pathways because many climatic variables have important
effects on conservation outcomes. Those variables include precipitation, tempera-
ture, wind, solar radiation, and atmospheric carbon dioxide, among others. Change
in any single variable also is complex (Fig. 3). A change in temperature, for exam-
ple, will affect conservation differently if that change primarily affects minimum,
maximum, or mean temperature. A change in a climatic variable also may differ
seasonally or geographically. The interaction between and among climatic variables
and conservation outcomes is dynamic and often nonlinear. Climatic variables inter-
act to magnify or dampen conservation effects. Likewise, conservation effects feed
back into the system and modify the influence of climatic variables. Those interac-
tions could have profound effects on soil, water, and related natural resources. Water
budgets, stream flow, and frequency and severity of floods and droughts may be
altered. Biotic communities, plant growth and development, and land use patterns
may shift. Those changes, in turn, may have important implications for soil, water,
and air quality, as well as fish and wildlife habitat.
As the main constituent of terrestrial ecosystem, the functions and processes

of soil changes in response to global climate change. Soil water reserve is one of
the main sources of water that can be utilized by vegetation. The potential change
of soil water induced by climate change may cause great change to ecological
environment and agricultural production. Globally, climate change affects average
temperatures and temperature extremes; timing and geographical patterns of
precipitation; snowmelt, runoff, evaporation, and soil moisture; the frequency of
disturbances such as drought, insect and disease outbreaks, severe storms and forest
fires; atmospheric composition and air quality; and patterns of human settlement
and land use change. Ecosystems and their services (land and water resources,
agriculture, biodiversity) experience a wide range of stresses, including pests and
pathogens, invasive species, air pollution, extreme events, wildfires and floods.
Climate change can cause or exacerbate direct stress through high temperatures,
reduced water availability, and altered frequency of extreme events and severe
storms. Understanding climate impacts on each of these sectors requires monitoring
many aspects of climate and a wide range of biological and physical responses.
Therefore, it could be concluded that a changing climate has effects on soil and
water resources in many ways. That means, climate change will affect soil and water
conservation through multiple pathways because many climatic variables have
important effects on conservation outcomes. Those variables include precipitation,
temperature, wind, solar radiation, and atmospheric carbon dioxide, among others.
3.4 Climate Change: A Blessing or a Curse for Agriculture?
Global climate change is a change in the long-term weather patterns that characterize
the regions of the world. Scientists state unequivocally that the earth is warming.
Natural climate variability alone cannot explain this trend. Human activities, espe-
cially the burning of coal and oil, have warmed the earth by dramatically increasing
the concentrations of heat-trapping gases in the atmosphere. The more of these
gases humans put into the atmosphere, the more the earth will warm in the decades
and centuries ahead. The impacts of warming can already be observed in many
places, from rising sea levels to melting snow and ice to changing weather patterns.
Climate change is already affecting ecosystems, freshwater supplies, and human
health. Although climate change cannot be avoided entirely, the most severe impacts
of climate change can be avoided by substantially reducing the amount of heat-
trapping gases released into the atmosphere. However, the time available for begin-
ning serious action to avoid severe global consequences is growing short. Global
Warming or climate change is a topic that increasingly occupies the attention of the
world (Table 2). Is it really happening? If so, how much of it is due to human activi-
ties? How far will it be possible to adapt to changes of climate? What action to
combat it can or should we take? How much will it cost? Or is it already too late for
useful action? Why carbon dioxide is both a blessing and a curse? Is climate change:
a blessing or a curse for agriculture?
Table 2 The impacts of climate change through both of the biophysical and socio-economical
effects. These different impacts include different agricultural issues from biophysical and socio-
economical point of view (Adapted from FAO 2007)
Biophysical effects Socio-economical effects
Physiological effects on crops, pasture, forests Decline in yields and production (exacer-
and livestock (quantity, quality); bating food insecurity);
Changes in land, soil and water resources Reduced marginal GDP from agriculture;
(quantity, quality);
Increased weed and pest challenges; Fluctuations in world market prices;
Shifts in spatial and temporal distribution of Changes in geographical distribution of
impacts; trade regimes;
Sea level rise, changes to ocean salinity; and Increased number of people at risk of
hunger and food insecurity; and
Sea temperature rise causing fish to inhabit Migration and civil unrest
different geographical ranges
GDP gross domestic production
Carbon dioxide might be a greenhouse gas, but it’s not necessarily bad for the
planet. Without it, there’d be no plant life and no human life as we know it. It’s only
toxic in high concentrations. And now the most important question is the climate
changing? This question could be answered as follows: It seems certain that the
world will be even more crowded and more connected. Will the increasing scale of
human activities affect the environment? In particular, will the world be warmer?
How is its climate likely to change?
Variations in day-to-day weather are occurring all the time; they are very much
part of our lives. The climate of a region is its average weather over a period that
may be a few months, a season or a few years. Variations in climate are also very
familiar to us. We describe summers as wet or dry, winters as mild, cold or stormy.
In many parts of the world, no season is the same as the last or indeed the same as
any previous season, nor will it be repeated in detail next time round. Most of these
variations we take for granted; they add a lot of interest to our lives. Those we par-
ticularly notice are the extreme situations and the climate disasters. Most of the
worst disasters in the world are, in fact, weather- or climate related.
Not all the climate changes will in the end be adverse. While some parts of the
world experience more frequent or more severe droughts, floods or significant sea
level rise, in other places crop yields may increase due to the fertilizing effect of
carbon dioxide. Other places, perhaps for instance in the sub-arctic, may become
more habitable. Even there, though, the likely rate of change will cause problems:
large damage to buildings will occur in regions of melting permafrost, and trees
in sub-arctic forests like trees elsewhere will not have time to adapt to new climatic
regimes. Scientists are confident about the fact of global warming and climate
change due to human activities. However, uncertainty remains about just how
large the warming will be and what will be the patterns of change in different parts
of the world. Although useful indications can be given, scientists cannot yet say
in precise detail which regions will be most affected. Intensive research is needed
to improve the confidence in scientific predictions (Houghton 2009).
Climate change is an increasingly urgent problem with potentially far-reaching

consequences for life on earth. Humans and wildlife are also exposed to an array of
chemical, physical, and biological stressors that arise largely from anthropogenic
activity, but also from natural sources. One of the consequences of climate change
that has recently attracted attention is its potential to alter the environmental distri-
bution and biological effects of chemical toxicants. There is growing awareness of
the importance of anticipating the effects of chemical pollution in the rapidly chang-
ing environment, and identifying and mitigating effects in those humans and eco-
systems most vulnerable (Noyes et al. 2009).
Today, global climate change is a fact. The climate has changed visibly, tangibly,
measurably. An additional increase in average temperatures is not only possible, but
very probable, while human intervention in the natural climate system plays an
important, if not decisive role (Porro 2002). Climate change is a major concern in
relation to the minerals sector and sustainable development. It is, potentially, one of
the greatest of all threats to the environment, to biodiversity and ultimately to our
quality of life (FTF 2002).
Climate on Earth has changed many times during the existence of our planet,
ranging from the ice ages to periods of warmth. During the last several decades
increases in average air temperatures have been reported and associated effects on
climate have been debated worldwide in a variety of forums. Due to its importance
around the globe, agriculture was one of the first sectors to be studied in terms of
potential impacts of climate change (Adams et al. 1990). According to studies car-
ried out by IPCC, average air temperatures will increase between 1.4°C and 5.8°C
by the end of this century, based upon modeling techniques that incorporated data
from ocean and atmospheric behavior. The possible impacts of this study, however,
are uncertain since processes such as heat, carbon, and radiation exchange among
different ecosystems are still under investigation. Less drastic estimates predict tem-
perature increase rates of 0.088°C per decade for this century (Kalnay and Cai
2003). Other investigators forecast for the near future that rising air temperature
could induce more frequent occurrence of extreme drought, flooding or heat waves
than in the past (Assad et al. 2004).
Studies of specific local impacts of climate changes have been conducted by
hundreds of research groups, many from organizations concerned with such matters
as seasonal crop forecasts, water supply and coastal protection. These groups have
found that climate change and sea-level rise of the magnitude and rates suggested
would greatly affect many natural systems like forests, rivers and wildlife, as well
as human activities and society. Examples include: (1) changes in natural productiv-
ity and biodiversity, with an increased rate of extinctions, (2) decreases in cereal
crop yields in most tropical and sub-tropical countries, and in temperate countries
for large warmings, (3) increased water shortages in many water-scarce regions due
to regional decreases in precipitation, increased evaporation and loss of glaciers and
seasonal snow storages, (4) adverse economic effects in many developing countries
for even small warmings, and for developed countries for larger warmings, (5) tens
of million of people on small islands and low-lying coastal areas at severe risk of
flooding from sea-level rise and storm surges, (6) increased threats to human health,
(7) increased inequities between poor and richer countries, (8) increased risk of
abrupt and irreversible climate changes (Pittock 2009).
In conclusion, climate change impacts are complex in that they can be both direct
and indirect. For example, more rain may lead directly to either greater or smaller
crop yields, depending on factors such as the type of crop, the soil and the present
climate. Indirect effects could include changes in supply and demand as a result of
these larger or smaller yields, both regionally and globally, and the resulting changes
in commodity prices, the profitability of farming, and the affordability of food and
effects on human health. Moreover, impacts can often be made more favorable by
changing strategies so as to minimize losses and maximize gains. It could be con-
cluded that climate change is a blessing and a curse for agriculture.
3.5 The Climate of Egypt
Egypt has an arid desert climate. It is hot or warm during the day, and cool at night.
In the coastal regions, temperature daytime temperatures range between an average
of minimum 14°C (57°F) in winter and average of maximum 30°C (86°F) in sum-
mer. In deserts the temperature varies to a great degree, especially in summer; it may
range from 7°C (44.6°F) at night, to 40°C (104.0°F) during the day. While the winter
temperatures in deserts do not fluctuate as wildly, they can be as low as 0°C (32°F)
at night, and as high as 18°C (64.4°F) during the day. Hot and dry Khamsin winds
blow in the Nile Delta region Egypt receives less than 80 mm (3.15 in) of precipita-
tion annually in most areas, although in the coastal areas it reaches 200 mm. It hardly
ever rains during the summer (Table 3; Zahran and Willis 2009).
Although Egypt is an arid country, its climate was wet in geological times.
The history of the climate in Egypt has been subject to many speculations based
on inference from geomorphological and archaeological studies: see for exam-
ple, Sandford (1934), Murray (1951) and Butzer (1959). Murray (1951) con-
cludes that regular rainfall ceased over Egypt below the 500 m contour some
time about the close of the Plio-Pleistocene period, three-quarters of a million
years ago and, though torrents from the Red Sea Hills have been able to maintain
their courses to the Nile through the foothills of the Eastern Desert, the Western
Desert has ever since been exposed to erosion by wind alone. The earlier European
glaciations seem to have left the Egyptian desert dry, but the long span of drought
was broken by at least two rainy interludes; the first when the deserts, both east
and west of the Nile, were habitable in Middle Palaeolithic times, the second,
with light rainfall, from about 8000–4000 BC. An occurrence of subsoil water
near the surface in the southern part of the Western Desert permitted people to
live there in oases till about 3000 BC when a drop of the water-table rendered
these places uninhabitable. The source of surface water all over the Eastern
Desert is the rainfall on the chains of the Red Sea Mountains. These, mountains
seem to intercept some orographic rain from the continental northerlies which
absorb their moisture through passage over the warm water of the Red Sea. The
62
Table 3 Comparison among the total greenhouse gas indicators (emissions of CO2, emissions per capita ratio and specific emission ratio) 1990/2000 in Egypt
(Adapted from UNDP 2011)
Population, GDP market price, Emissions, Emission ton Emissions per capita Emission ton CO2e Specific emission
Year million billion US$ million ton CO2e CO2e per capita ratio (2000/1990), % per thousand US$ ratio (2000/1990), %
1990 52.6 35.16 116.6 2.2 – 3.3 –
2000 63.3 99.74 193.3 3.1 137% 1.9 58
GDP gross domestic production, US$ United States dollar
H.R. El-Ramady et al.
mountain rains may feed the wadis of the Eastern Desert with considerable tor-
rential flows (Zahran and Willis 2009).
According to Ayyad and Ghabbour (1986), Egypt can be divided into two hyper
arid and two arid provinces as follows:
1. Hyper arid provinces:
(a) Hyper arid with a mild winter, where mean temperature of the coldest month
between 10°C and 20°C and a very hot summer with mean temperature of
the hottest month more than 30°C, including the southwestern part of the
Western Desert.
(b) Hyper arid with a mild winter and a hot summer, i.e. mean temperature of
the hottest month 20–30°C covering the Eastern Desert and the northeastern
part of the Western Desert and Gebel Uweinat area.
2. Arid provinces:
(a) The northern section with winter rainfall which extends along the
Mediterranean coast and the Gulf of Suez. This section is divided into two
provinces by the UNESCO/FAO map of 1963: the coastal belt province
under the maritime influence of the Mediterranean, with a shorter dry period
attenuated, and a more inland province with a longer dry period accentuated
and an annual rainfall of 20–100 mm. Both provinces are characterized by a
mild winter and a hot summer.
(b) A southern section with winter rainfall which includes one province – the
Gebel Elba area of the Red Sea coast of Egypt (Zahran and Willis 2009).
Therefore, Egypt has an arid desert climate. It is hot or warm during the day, and
cool at night. In the coastal regions, temperature daytime temperatures range between
an average of minimum 14°C in winter and average of maximum 30°C in summer.
In deserts the temperature vary to a great degree, especially in summer; they may
range from 7°C at night, to 40°C during the day. While the winter temperatures in
deserts do not fluctuate as wildly, they can be as low as 0°C at night, and as high as
18°C during the day. Hot and dry Khamsin winds blow in the Nile Delta region.
Egypt receives less than 80 mm of precipitation annually in most areas, although in
the coastal areas it reaches 200 mm. It hardly ever rains during the summer. It could
be also divided the climate of Egypt to hyper arid provinces and arid provinces.
3.6 Situation Analysis for Egypt and Climate Changes
Situation analysis of Egypt and climate changes has been reported as follows (UNDP
2011):
1. In 1994, Egypt ratified the United Nations Framework Convention on Climate
Change (UNFCCC). The Intergovernmental Panel on Climate Change in its
Third assessment report, IPCC (1995), identified Egypt’s Mediterranean coast

and the Nile Delta as vulnerable regions to sea level rise. In this respect, Egypt
set up the Climate Change institutional structure at the national level, and a
Climate Change Unit was established in 1996 in Egyptian Environmental Affairs
Agency.
2. In 2007, Egyptian Prime Minister issued Decree No. 272 to reform the National
Climate Change Committee that was established in 1997. The new Climate
Change Committee is chaired by the Minister of State for Environmental Affairs
and includes members representing a wide range of governmental and nongov-
ernmental representatives. In addition, Ministry of State for Environmental
Affairs upgraded Climate Change Unit to be a Central Department in Egyptian
Environmental Affairs Agency in 2009, in order to strengthen the climate change
institutional structure on the national level.
3. Two ministerial climate change committees in the Ministry of Agriculture &
Land Reclamation and the Ministry of Irrigation & Water Resources have been
established. In addition, a climate change information centre in the Agriculture
Research Centre has been established. However, many barriers still exist that are
challenging Egypt efforts to comply with United Nations Framework Convention
on Climate Change such as inadequate capacity and weak coordination and
cooperation between governmental bodies, Non-governmental Organizations
and private sector. Furthermore there is a lack of mainstreaming the adaptation
measures in the national planning process, particularly in comparison with miti-
gation measures.
4. Egypt submitted Initial and Second National Communication reports to United
Nations Framework Convention on Climate Change in 1999 and 2010, respec-
tively. According to Initial National Communication and Second National
Communication Egypt’s most vulnerable sectors to climate change are identified
as follows: (1) coastal zones, (2) water resources and (3) agriculture. The sea
level rise is the cause of the most serious climate change impacts that threatens
the densely populated River Nile Delta which includes extensive infrastructure
and fertile agriculture lands. In this respect, sea level rise is expected to inundate
large areas of low lying lands in the Nile Delta and sea water intrusion will
increase water logging conditions and soil salinity in other lands. Furthermore,
there is also a high degree of uncertainty regarding the climate change impacts
on the annual Nile flood, the expected decline in precipitation along the North
Coast and a projected increase in the population estimated between 115 and 179
million by 2050. Moreover, temperature rise is expected to reduce the productiv-
ity of major crops, increase crop water requirements coupled with an expected
water stress and loss of some lands and fertility in the Nile Delta and conse-
quently the overall food production may be significantly reduced. Impacts of
climate change on other vulnerable sectors to climate change will be further
investigated in the Third National Communication. Accordingly, climate change
risks may threaten Egypt’s efforts to achieve the Millennium Development Goals
and to face those threats; Initial National Communication and Second National
Communication presented several adaptation measures to climate change

impacts, as well as, many mitigation measures to play an effective role in achiev-
ing the main target of the United Nations Framework Convention on Climate
Change.
Review information on vulnerability and impacts provided in the Second National
Communication and re-confirm gaps in data collection and analysis for all areas,
with special emphasis on the following four priority areas identified under the
Second National Communication and designated by the Government as continuing
priority areas for the Third National Communication:
• Agriculture updating data and analysis on cereal crops production, fibre crops;
and livestock, as well as, study other new areas such as insects, plant disease and
fisheries
• Water resources particularly new data and resulting analysis from the Regional
Circulation Model that developed under Climate Change Risk Management
Program
• Coastal Zones updating all related information to Sea Level Rise and its impacts
on Costal areas in Egypt and particularly in Delta
• Studying new areas that were not covered in Second National Communication
such as biodiversity. As well as, give more attention to health, human habitat &
settlement and tourism that were not addressed adequately in Second National
Communication (UNDP 2011).
From the previous, it could be concluded that the situation analysis of Egypt and
climate changes started from ratification of Egypt to the United Nations Framework
Convention on Climate Change in 1994, establishment of Climate Change Unit in
1996 in the Egyptian Environmental Affairs Agency, issue the Egyptian Prime
Minister Decree No. 272 to reform the National Climate Change Committee in
2007, establishment of two ministerial climate change committees in the Ministry
of Agriculture & Land Reclamation and the Ministry of Irrigation & Water
Resources, and submission Egypt to Initial and Second National Communication
reports to United Nations Framework Convention on Climate Change in 1999 and
2010, respectively.
4 Climate Changes and Agriculture in Egypt
The natural greenhouse effect raises the temperature of the planet to 33°C, thus mak-
ing it habitable. On average, 343 W m−2 of sunlight fall on the earth, roughly 1/3 of
which is reflected back into space. The other 2/3 reaches the ground, which re-radiates
it as longer wavelength, infrared radiation. Some of this is blocked by greenhouse
gases, thereby warming the atmosphere. Naturally occurring greenhouse gases include
water vapor, CO2, methane (CH4) and nitrous oxide (N2O). Reducing emissions
of CO2 could be achieved by switching to renewable energy (IPCC 1996).
Nature provides freshwater through the hydrologic cycle. The process is as

follows: production of vapors above the surface of the liquids, the transport of
vapors by winds, the cooling of air–vapor mixture, condensation and precipitation
(Salem 2012).
In fact, climate is a primary determinant of agricultural productivity. In turn, food
and fiber production is essential for sustaining and enhancing human welfare. Hence,
agriculture has been a major concern in the discussions on climate change. Food sup-
ply vulnerability to climate change is an issue in two different ways. First, future
food supply may be directly threatened by climate change. Second, food supply
capacity may be altered by efforts to reduce greenhouse gases emissions as society
tries to mitigate future implications of climate change. Agronomic and economic
impacts from climate change depend primarily on two factors: (1) the rate and
magnitude of change in climate attributes and the agricultural effects of these changes,
and (2) the ability of agricultural production to adapt to changing environmental
conditions (McCarthy et al. 2001).
Climate is the single most important determinant of agricultural productivity,
primarily through its effects on temperature and water regimes. For example, the
physiographic boundaries of principal biomes are determined by mean annual tem-
perature and soil water regimes. Climate change is therefore expected to alter the
biophysical environment of growing crops and to influence biomass productivity
and agronomic yields (Rosenzweig and Hillel 1998).
Positive effects may be associated with the fertilization effects of CO2 enrich-
ment, increases in the duration of growing seasons in higher latitudes and mountain
ecosystems, and possible increase in soil water availability in regions with an
increase in annual precipitation. Each 1°C increase in temperature may lead to a
10-day increase in the growing season in northern Europe and Canada. The CO2
fertilization effect is real. However, the net positive effect may be moderated by
other factors, such as the effective rooting depth and nutrient availability. Further,
the productivity per unit of available water is expected to rise by 20–40% (van de
Geijn and Goudriaan 1996).
Negative effects of projected climate change on agriculture may be due to
increases in respiration rate as temperature rises with attendant decreases in net
primary productivity (NPP); increases in the incidence of pests and diseases; short-
ening of the growing period in some areas; decrease in water availability as rainfall
patterns change; poor vernalization; and increased risks of soil degradation caused
by erosion and possible decline in SOC concentration. The yield of rice has been
estimated to decrease by 9% for each 1°C increase in temperature. Phillips et al.
(1996), using the explicit planetary isentropic coordinate (EPIC) model to examine
the sensitivity of corn and soybean yields to climate change, projected a 3% decrease
in both corn and soybean yields in response to a 2°C increase in temperature from a
baseline precipitation level. However, a 10% precipitation increase balanced the
negative effect of a 2°C temperature increase. The effects of climate change on crop
yields may be more negative at lower latitudes and generally positive at middle and
high-middle latitudes. Further, crop growth is more affected by extremes of weather
than by averages. The annual average changes in temperature or precipitation used
in most predictive models do not reflect the short-term effects of so-called extreme
events — droughts, floods, freezes, or heat waves (Lal 2005).
4.1 Climate Changes and Its Impact on Agriculture
Driven mainly by population and economic growth, total world food consumption
is expected to increase over 50% by 2030 and may double by 2050 (Barker et al.
2007). Most of the increase in food production in the next decades is expected to
occur through further intensification of current cropping systems rather than through
opening of new land into agricultural production. Intensification of cropping sys-
tems has been a highly successful strategy for increasing food production. The best
example is the well-known success of the Green Revolution, where the adoption of
modern varieties, irrigation, fertilizers and agrochemicals resulted in dramatic
increases in food production. However, this strategy also resulted in unexpected
environmental consequences, one of them being the emissions of greenhouse gases
into the atmosphere. Therefore, future strategies that promote further intensification
of agriculture should aim at the development of sustainable cropping systems that
not only consider increasing food production but that also look at minimizing envi-
ronmental impact (Ortiz-Monasterio et al. 2010).
At present, 40% of the Earth’s land surface is managed for cropland and pasture
(Foley et al. 2005). The most important cropping systems globally, in terms of meet-
ing future food demand, are those based on the staple crops, rice, wheat and maize.
Rice and maize are each grown on more than 155 million ha (FAOSTAT 2009). In
addition, rice is the staple food of the largest number of people on Earth. The geo-
graphic distribution of rice production gives particular significance to Asia where
90% of the world’s rice is produced and consumed. Maize is produced mainly in the
Americas, followed by Asia and then Africa. Maize is important as a staple crop
(mainly in developing countries) but it is also important as animal feed and, increas-
ingly, as biofuel. Wheat is the most widely grown crop, covering more than 215
million ha around the world, with Asia covering close to 50% of the world wheat
area (FAOSTAT 2009).
Without additional policies, agricultural N2O and CH4 emissions are projected to
increase by 35–60% and ~ 60%, respectively, to 2030, thus increasing more rapidly
than the 14% increase of non-CO2 greenhouse gases observed from 1990 to 2005
(Barker et al. 2007). Improved agricultural management enhances resource-use
efficiencies, often reducing emissions of more than one greenhouse gas. The effec-
tiveness of these practices depends on factors such as climate, soil type and farming
system. About 90% of the total mitigation arises from sink enhancement (soil C
sequestration) and about 10% from emission reduction. In spite of inherent uncer-
tainties in such estimates, it can be concluded that the topic of this review, which
addresses the second option (improved cropland management) and the fifth option
(improved rice management), comprises a sizable portion of the overall mitigation
potential of agriculture (Ortiz-Monasterio et al. 2010).
Promoting agricultural practices that mitigate climate change by reducing GHG
emissions is important, but those same practices also have to improve farmer pro-
duction and income and buffer the production system against the effects of changes
in climate. The overall impact predicted by climate change models vary but we are
now locked into global warming and inevitable changes to climatic pattern that are
likely to exacerbate existing rainfall variability and further increase the frequency of
Fig. 4 Points that must be considered while doing the study of impacts of climatic changes on agri-
culture, which include soil, water, plant and the agroecosystem (Adapted from Khan et al. 2009)
climatic extremes. Where excess rain occurs, extreme rainfall events will increase
leading to flooding and soil erosion. In low rainfall, drought-prone areas there is
general acceptance in the science community of more frequent moisture stress
because of failed rainfall patterns and increased evaporation caused by higher tem-
peratures (Cooper et al. 2008). In Africa specifically, the projected combined
impacts of climate change and population growth suggest an alarming increase in
water scarcity for many countries, with 22 of the 28 countries considered likely to
face water scarcity or water stress by 2025. This in turn will curtail the ability of
irrigated agriculture to respond to the expanding food requirements of tomorrow’s
Africa (Rosegrant et al. 2002).
In order to cope with the increased climate risk, agricultural systems will have to
be more robust and resilient to buffer for extreme weather events such as drought,
flooding, etc. (Fig. 4). It is paramount that new agricultural practices not only pre-
vent further soil degradation but also improve system resilience through increased
soil organic matter, improved water-use efficiency as well as nutrient-use efficiency,
and increased flora and fauna biodiversity. However, the management of agriculture
to cope with greenhouse gases emissions and the negative effects of climate change
on food production lies in the hands of farmers, pastoralists and forest managers
whose decisions are determined by multiple goals (Hobbs and Govaerts 2010).
The impacts of climate change on agriculture are expected to be widespread
across the globe, although studies suggest that African agriculture is likely to be
most affected due to heavy reliance on low-input rainfed agriculture and due to its
low adaptive capacity (Mertz et al. 2009). Broadly speaking, climate change is
likely to impact crop productivity directly through changes in the growing environ-
ment, but also indirectly through shifts in the geography and prevalence of agricul-
tural pests and diseases, associated impacts on soil fertility and biological function,
and associated agricultural biodiversity. While many impact predictions tend
towards the negative, increased CO2 will also contribute to enhanced fertilization –
although there is significant debate as to the extent to which this may increase plant
growth. This section looks at these issues, concentrating entirely on the expected
biophysical impacts (Jarvis et al. 2010).
Climate change due to anthropogenically-generated greenhouse gases and aero-
sols has been recognized as a serious threat to the earth’s ecosystems and its inhabit-
ants, and the dangers associated with climate change will increase in severity in
coming decades in the absence of measures to curb the production of the responsi-
ble pollutants e.g., carbon dioxide, methane, nitrous oxide. Numerous scientific
articles and peer-reviewed reports have demonstrated the current and potential
future effects of the climate change that result from these increased greenhouse gas
emissions (IPCC 2007).
Among the resource sectors that received early attention regarding possible cli-
mate change effects has been agriculture and it has continued to receive consider-
able attention since that early research. Work in this area has become more
sophisticated over time and is now connected explicitly to estimates of economics
in the agricultural sector (Reilly 2010) and risk of hunger. As is the case with many
impact areas, studies of possible adaptation to climate change have come to the fore
and become increasingly important (Easterling 2010), and this is particularly strik-
ing for agriculture where studies of adaptation to climate change appeared early in
the history of climate change research. The importance of adaptation studies has
also put more emphasis on the need for more detailed information regarding future
regional climate change (Mearns 2011).
It could be concluded that climate change and agriculture are interrelated pro-
cesses, both of which take place on a global scale. At the same time, agriculture has
been shown to produce significant effects on climate change, primarily through the
production and release of greenhouse gases such as CO2, CH4, and nitrous oxide,
but also by altering the Earth’s land cover, which can change its ability to absorb or
reflect heat and light, thus contributing to radiative forcing. Land use change such
as deforestation and desertification, together with use of fossil fuels, are the major
anthropogenic sources of CO2; agriculture itself is the major contributor to increas-
ing CH4 and nitrous oxide concentrations in Earth’s atmosphere. Agronomic and
economic impacts from climate change depend primarily on two factors: (1) the rate and
magnitude of change in climate attributes and the agricultural effects of these changes,
and (2) the ability of agricultural production to adapt to changing environmental
conditions. Temperature, precipitation, atmospheric carbon dioxide content, the

incidence of extreme events and sea level rise are the main climate change related
drivers which impact agricultural production. Briefly, the main categories of agri-
cultural productivity implications are: crops and forage productivity and production
cost, soil suitability for agricultural production, livestock productivity and produc-
tion cost and irrigation water supply.
4.2 Climate Change’s Impacts on Global Crop Productivity
The major concerns for crop productivity as a result of increased levels of green-
house gases are related to warmer temperatures and altered amounts and patterns of
rainfall. Both average temperature and temperature variability are predicted to
increase. Average global temperatures are predicted to increase by 0.6–2.5°C over
the next 50 years with significant spatial variation. While this will permit cultivation
of crops in areas of the world which are currently too cold e.g. Siberia and northern
America and extend the potential growing season for others, it will also threaten the
viability of crops in many of the major areas of production. Simulation models sug-
gest that wheat yields in south-east Australia may decrease by about 29% (Anwar
et al. 2007) and direct studies in the Philippines have shown that irrigated rice yields
decrease by 10% for each 1°C increase in the minimum night-time temperature
although the maximum temperature has no effect (Peng et al. 2004).
Higher temperatures will shorten the life cycle of most crops, by accelerating
development and hastening senescence, thereby decreasing the time available to
harvest light and produce biomass. The effects on phenology vary both between
species and with environment. Perennial crops may respond more strongly to an
increased temperature than annual crops (Estrella et al. 2007). Other effects such as
drought or an increase in ozone concentrations can exacerbate these effects. The
decreased time available to harvest light and produce biomass contributes to yield
reductions at elevated temperatures (Parry and Hawkesford 2010).
Our knowledge of water use is as poor as our knowledge of water resources per-
haps poorer. Information is largely incomplete particularly for agriculture, the larg-
est user and is lacking altogether for some countries. Only limited disaggregated
information exists, and even this shows deficiencies of validity and homogeneity
and provides extremely poor information on trends. The quality of information
systems varies with each country, but there are common difficulties: (1) Statistics on
the magnitude of demand and withdrawal are often estimated rather than based on
data that are measured or collected from censuses. The level of uncertainty varies,
but is particularly high for agriculture. (2) Sectors of use are not defined homoge-
neously and are not well disaggregated. (3) Adequate historical datasets are rare,
and the dates of available statistics are not always explicit. (4) Lack of agreed
terminology leads to discrepancies in data compilation and analyses. Agriculture
is by far the main user of water. Irrigated agriculture accounts for 70% of water
withdrawals, which can rise to more than 80% in some regions. Although increasing
in urbanized economies, industrial including energy use accounts for only 20% of
total water use and domestic use for about 10%. Water withdrawals for energy
generation, hydropower and thermo-cooling are on the rise, but energy is one of
the economic sectors that consumes the least water and it returns most of the water
withdrawn back to the water system (about 95%). This is only a partial picture of
sectoral usage as there are many unaccounted-for uses. Little is known about water
use in informal urban settlements or informal irrigation systems, both of which are
generally unaccounted for in official statistics (Connor et al. 2009).
Nowadays, climate changes, their causes and consequences, gained importance
in many other areas of interest for sustainable life on Earth. The subject is, however,
controversial. Understanding how climate changes will impact mankind in the
decades to come is of paramount importance for our survival. Temperature, carbon
dioxide (CO2) and ozone (O3) directly and indirectly affect the production and
quality of fruit and vegetable crops grown in different climates around the world.
Temperature variation can directly affect crop photosynthesis, and a rise in global
temperatures can be expected to have significant impact on postharvest quality by
altering important quality parameters such as synthesis of sugars, organic acids,
antioxidant compounds and firmness. Rising levels of CO2 also contribute to global
warming, by entrapping heat in the atmosphere. Prolonged exposure to concentra-
tions could induce higher incidences of tuber malformation and increased levels of
sugars in potato and diminished protein and mineral contents, leading to loss of
nutritional and sensory quality. Increased levels of O3 in the atmosphere can lead to
detrimental effects on postharvest quality of fruit and vegetable crops. Elevated
levels of O3 can induce visual injury and physiological disorders in different species,
as well as significant changes in dry matter, reducing sugars, citric and malic acid,
among other important quality parameters (Moretti et al. 2010).
Besides increase in temperature and its associated effects, climate changes are
also a consequence of alterations in the composition of gaseous constituents in the
atmosphere. CO2, also known as the most important greenhouse gas, and O3 con-
centrations in the atmosphere are changing during the last decade and are affecting
many aspects of fruit and vegetable crops production around the globe (Felzer
et al. 2007).
Exposure to elevated temperatures can cause morphological, anatomical, physi-
ological, and, ultimately, biochemical changes in plant tissues and, as a conse-
quence, can affect growth and development of different plant organs. These events
can cause drastic reductions in commercial yield. However, by understanding plant
tissues physiological responses to high temperatures, mechanisms of heat tolerances
and possible strategies to improve yield, it is possible to predict reactions that will
take place in the different steps of fruit and vegetable crops production, harvest and
postharvest (Kays 1997). Temperature increase and the effects of greenhouse gases
are among the most important issues associated with climate change. Studies have
shown that the production and quality of fresh fruit and vegetable crops can be
directly and indirectly affected by high temperatures and exposure to elevated levels
of carbon dioxide and ozone. Temperature increase affects photosynthesis directly,
causing alterations in sugars, organic acids, and flavonoids contents, firmness and
antioxidant activity. Higher temperatures can increase the capacity of air to absorb
water vapor and, consequently, generate a higher demand for water. Higher evapo-
transpiration indices could lower or deplete the water reservoir in soils, creating
water stress in plants during dry seasons. For example, water stress is of great con-
cern in fruit production, because trees are not irrigated in many production areas
around the world. It is well documented that water stress not only reduces crop
productivity but also tends to accelerate fruit ripening (Henson 2008).
Carbon dioxide concentrations are increasing in the atmosphere during the last
decades (Mearns 2000). The current atmospheric CO2 concentration is higher than
at any time in the past 420,000 years (Petit et al. 1999). Further increases due to
anthropogenic activities have been predicted. Carbon dioxide concentrations are
expected to be 100% higher in 2100 than the one observed at the pre-industrial era
(IPCC 2007). Ozone concentration in the atmosphere is also increasing. Even low-
levels of ozone in the vicinities of big cities can cause visible injuries to plant tissues
as well as physiological alterations (Felzer et al. 2007). Carbon dioxide accumula-
tion in the atmosphere has directly effects on postharvest quality causing tuber mal-
formation, occurrence of common scab, and changes in reducing sugars contents on
potatoes. High concentrations of atmospheric ozone can potentially cause reduction
in the photosynthetic process, growth and biomass accumulation. Ozone-enriched
atmospheres increased vitamin C content and decreased emissions of volatile esters
on strawberries. Tomatoes exposed to ozone concentrations ranging from 0.005 to
1.0 m mol mol−1 had a transient increase in b-carotene, lutein and lycopene
contents.
IPCC (2007) concluded that ‘in mid- to high latitude regions, moderate warming
benefits crop and pasture yields, but even slight warming decreases yields in season-
ally dry and low-latitude regions, i.e. medium confidence’. In IPCC language, mod-
erate warming is in the range of 1–3°C. Smallholder and subsistence farmers,
pastoralists and artisanal fisher-folk will suffer complex, localized impacts of cli-
mate change, i.e. high confidence. Food and forestry trades are projected to increase
in response to climate change with increased dependence on food imports for most
developing countries, i.e. medium to low confidence. The report further concluded
that warming beyond 2–3°C was likely to result in yield declines in all areas. This
analysis was based on a synthesis of 69 studies, which was a vast improvement on
the handful of studies used in the Third Assessment Report, AR3 (IPCC 2001).
But even since the IPCC, AR4 (2007) there has been a much larger number of
studies which examine the impacts of climate change on crop production and
yields, including global multi-crop studies, down to regional and national studies
on individual crops. This chapter summarizes the IPCC findings, and provides a
more detailed analysis of impact studies arising from 2006 to 2009. There are
fairly consistent pictures drawn by different studies that show the potential effects
of changing climates (Lobell et al. 2008). These all show steeply increasing trends
in adverse impacts, particularly in food insecure regions among the tropics, which
are likely to increase the extent to which these regions are food insecure, especially
taking into account that most of these regions present the least adaptive capacity
(Jarvis et al. 2010).
Grain yields are expected to fall in developing countries; however, the opposite
is likely to happen in developed countries (IPCC 2007). Geographies of changes
may influence yield responses: in high latitudes, where most of the developed coun-
tries are located, increased temperatures could increase the duration of growing
seasons, thus benefiting farmers. However, in developing countries, which are
mostly located in the tropics, this effect would not be observed. Investment capacity
within the different agricultural sectors needs to be considered if yield losses are to
be offset. Moreover, yield reductions will certainly result in increases in prices of
agricultural goods, and this impact will be greater for food insecure regions (Jarvis
et al. 2010).
Therefore, the major concerns for crop productivity as a result of increased levels
of greenhouse gases are related to warmer temperatures and altered amounts and
patterns of rainfall. Higher temperatures will shorten the life cycle of most crops, by
accelerating development and hastening senescence, thereby decreasing the time
available to harvest light and produce biomass. The effects on phenology vary both
between species and with environment and perennial crops may respond more
strongly to an increased temperature than annual crops. Other effects such as drought
or an increase in ozone concentrations can exacerbate these effects. The decreased
time available to harvest light and produce biomass contributes to yield reductions
at elevated temperatures.
4.3 Impacts of Climate Changes on Crop Physiology
Agriculture accounts for 70% of freshwater withdrawals from rivers, lakes and
aquifers up to more than 90% in some developing countries. Furthermore, unlike in
industrial and domestic uses, where most of the water returns to rivers after use, in
agriculture a large part of water is consumed by evapotranspiration. Many irrigation
systems, however, return a large amount of water to the system after use. Biomass
cannot be produced without water. The source of all food is photosynthesis. Biomass
is processed through the food chain, which describes the flow of energy and feeding
relationship between species: from primary producers, i.e. plants to herbivores to
carnivores. It could be estimated how much water is needed to sustain our diets by
calculating the water lost in evapotranspiration based on crop physiology. Depending
on local climate, varieties and agronomical practices, it takes 400–2,000 l of evapo-
transpiration daily to produce 1 kilogram (kg) of wheat, and 1,000–20,000 l per
kilogram of meat, depending on the type of animal, feed and management practices.
Based on these values, researchers have estimated daily water requirements to sup-
port diets, ranging from 2,000 to 5,000 l of water per person per day. FAO uses
2,800 kilocalories (kcal) per person at the national level as a threshold for food
security. As a rule of thumb, it can therefore be estimated that 1 l of water is needed
to produce 1 kcal of food. Because of the low energy efficiency of the food chain,
protein-rich diets require substantially more water than vegetarian diets (Connor
et al. 2009).
Over the past 800,000 years, atmospheric (CO2) changed between 180 ppm (gla-
cial periods) and 280 ppm (interglacial periods) as Earth moved between ice ages.
From pre-industrial levels of 280 ppm, (CO2) has increased steadily to 384 ppm in
2009, and mean temperature has increased by 0.76°C over the same time period.
Projections to the end of this century suggest that atmospheric (CO2) will top
700 ppm or more, whereas global temperature will increase by 1.8–4.0°C, depend-
ing on the greenhouse emission scenario (IPCC 2007). There is growing evidence
suggesting that many crops, notably C3 crops, may respond positively to increased
atmospheric CO2 in the absence of other stressful conditions (Long et al. 2004), but
the beneficial direct impact of elevated CO2 can be offset by other effects of climate
change, such as elevated temperatures, higher tropospheric ozone concentrations
and altered patterns of precipitation (Easterling et al. 2007; Da Matta et al. 2010).
It is now universally accepted that increased atmospheric concentrations of
‘greenhouse gases’ are the main cause of the ongoing climate change (Forster et al.
2007) and that these changes are expected to have important effects on different
economic sectors, e.g. agriculture, forestry, energy consumptions, tourism, etc.
(Hanson et al. 2007). Since agricultural practices are climate-dependent and yields
vary from year to year depending on climate variability, the agricultural sector is
particularly exposed to changes in climate. In Europe, the present climatic trend
indicates that in the northern areas, climate change may primarily have positive
effects through increases in productivity and in the range of species grown (Alcamo
et al. 2007), while in southern areas (i.e. the Mediterranean basin) the disadvantages
will predominate with lower harvestable yields, higher yield variability and a reduc-
tion in suitable areas for traditional crops (Moriondo et al. 2010).
For climate change impact assessment, crop growth models have been widely
used to evaluate crop responses, i. e. development, growth and yield by combining
future climate conditions, obtained from General or Regional Circulation Models,
with the simulation of CO2 physiological effects, derived from crop experiments
(Ainsworth and Long 2005). Many of these impact studies were aimed at assessing
crop development shifts and yield variations under changes in mean climate condi-
tions. These analyses showed that increasing temperatures generally shortened the
growing period of commercial crops (Giannakopoulos et al. 2009), resulting in a
shorter time for biomass accumulation. On the other hand, changes in yields were
not homogeneous and dependent on crop phenology, e.g. summer and winter crops,
crop type, e.g. C3 and C4 plants or environmental conditions, e. g. water and nutrient
availability (Moriondo et al. 2010).
Other studies stressed that changes in climate variability, as can be expected in a
warmer climate, may have a more profound effect on yield than changes in mean
climate (Porter and Semenov 2005). As such, policy analysis should not rely on
scenarios of future climate involving only changes in means. Furthermore, the
changes in the frequency of extreme climatic events during the more sensitive
growth stages have been recognized as a major yield-determining factor for some
regions in the future (Easterling and Apps 2005; Schneider et al. 2007). Temperatures
outside the range of those typically expected during the growing season may have
severe consequences on crops, and when occurring during key development stages
they may have a dramatic impact on final production, even in case of generally
favorable weather conditions for the rest of the growing season. Many studies high-
lighted the potential of heat stresses during the anthesis stage as a yield reducing
factor (Challinor et al. 2005), while others pointed out that the joint probability of
heat stress-anthesis is likely to increase in future scenarios (Alcamo et al. 2007).
Accordingly, both changes in mean climate and climate variability including
extreme events should be considered for a reliable climate change impact assess-
ment in agriculture. An example is the summer heat wave of 2003 (Schaer et al.
2004), taken as an indicator of the future climate change, which reduced cereal
production in Europe by 23 MT with respect to 2002. The reason for this reduction
was attributed to the shorter growing season combined with a higher frequency of
extreme events, both in terms of maximum temperatures and longer dry spells
(Olesen and Bindi 2004). In contrast, climate change impact assessments carried
out so far have not included direct simulations of heat stress impact on crop yield
(Schneider et al. 2007) resulting in a probable underestimation of yield losses
(Moriondo et al. 2010).
The temperature response of crop growth and yield must be considered to predict
the (CO2) effects. The threshold developmental responses of crops to temperature
are often well defined, changing direction over a narrow temperature (Porter and
Semenov 2005). High temperatures reduce the net carbon gain in C3 species by
increasing photorespiration; by reducing photorespiration, (CO2) enrichment is
expected to increase photosynthesis more at high than at low temperatures, and thus
at least partially offsetting the temperature effects of supra-optimal temperatures on
yield. Therefore, yield increases at high (CO2) should occur most frequently in
regions where temperatures approximate the optimum for crop growth. Conversely,
in regions where high temperatures already are severely limiting, further increases
in temperature will depress crop yield regardless of changes in (CO2) (Polley 2002).
In fact, results of mathematical modeling suggest that, in mid- to high-latitude
regions, moderate to medium local increases in temperature (1–3°C), along with
associated CO2 increase and rainfall changes, can have beneficial impacts on crop
yields, but in low-latitude regions even moderate temperature increases (1–2°C) are
likely to have negative impacts on yield of major cereals (Easterling et al. 2007).
Thus, climate change may impair food production, particularly in developing coun-
tries, most of which are located in tropical regions with warmer baseline climates
(Da Matta et al. 2010).
In addition to crop growth and yield, crop quality is also expected to be affected
by global climatic changes. Crop quality is thought to be a multi-faceted and com-
plex subject involving growth, assimilate partitioning and storage, and pre- and
post-harvest, including nutritional, technological and environmental facets (Hay
and Porter 2006). Elemental, e.g. zinc, iodine and macromolecular, e.g. protein
composition in plant tissues are expected to change in a future high-CO2 world
(Taub et al. 2008). In this context, crop physiologists will need to take more account
of the interests of breeders and processors by studying, quantifying and modeling
the differences not only in increasing yields but also in food quality among crop
varieties and species in climate change scenarios (Hay and Porter 2006). The efforts
to understand the impact of elevated (CO2), temperature and other ongoing climatic
changes on food crops are crucial to estimate food production in the future. The
present review, which is by no means exhaustive, is mainly focused on the current
understanding of the consequences of climatic changes (mainly CO2 enrichment
and temperature) on crop physiology and chemistry (Da Matta et al. 2010).
Crops sense and respond directly to rising (CO2) through photosynthesis and
stomatal conductance, and this is the basis for the CO2 fertilization effect on crop
yield (Long et al. 2006). These responses are highly dependent on temperature
(Polley 2002). Therefore, understanding how crop species will respond to these
environmental changes is crucial for maximizing the potential benefits of elevated
CO2, for which agronomic practice needs to adapt as both temperature and CO2 rise
(Challinor and Wheeler 2008). In fact, both chemical and microbiological risks are
foreseen to impair food and feed safety as a consequence of climate change: in par-
ticular, mycotoxins, pesticide residues, trace metals and other chemicals could
affect food and feed safety (Miraglia et al. 2009). There is, therefore, an urgent need
for scientific research that can improve our understanding of the interactions of ris-
ing atmospheric (CO2) with other environmental variables, such as temperature,
water supply and ozone concentration, as well as with biotic factors such as pests
and diseases, under real field conditions. In doing so, it is necessary not only to
quantify the effects of climatic changes on crop production but also on food quality.
It is also necessary to assess responses of crops other than the key cereal grains, and
in climate regions other than temperate ones, notably those of importance to devel-
oping countries in the tropics and subtropics (Tubiello et al. 2007). Furthermore,
since distinct varieties seem to respond differently to elevated CO2 and temperature
in terms of harvestable yield, future research should be also directed towards select-
ing promising genotypes for a changing global climate (Da Matta et al. 2010).
From previous, it could be concluded that now universally is accepted that
increased atmospheric concentrations of greenhouse gases are the main cause of the
ongoing climate change and that these changes are expected to have important
effects on different economic sectors, e.g. agriculture, forestry, energy consump-
tions, tourism, etc. The increasing temperatures generally shortened the growing
period of commercial crops, resulting in a shorter time for biomass accumulation.
The changes in yields are not homogeneous and dependent on crop phenology, e.g.
summer and winter crops, crop type, e.g. C3 and C4 plants or environmental condi-
tions, e. g. water and nutrient availability.
4.4 Effects of Climatic and Global Change on Water Scarcity
Egypt has already reached the water poverty limit and needs a much greater share
of Nile water in year 2050 to cover the shortage. Surface freshwater pollution has
embarked on a critical path. One climate change scenario predicts that the Nile dis-
charge may decrease to 3/4 of its present volume if CO2 emissions double. Low cost solar
water desalination is a strategic solution for Egypt. The number of desalination
plants has increased in the last 30 years and generated 2333.963 m3 day−1 in 2004
(SRU 2006). There is a trend in Egypt to apply desalination to meet the require-
ments of industry, tourism, petroleum, electricity, health and reconstruction.
The desalination plants are located on the Red Sea coast, in south Sinai and on the
northern coast (Salim 2012).
Climate change projections also indicate an increased likelihood of droughts and
variability of precipitation – in time, space, and intensity – that would directly
influence water resources availability. The combination of long-term change, e. g.
warmer average temperatures and greater extremes, e. g. droughts can have decisive
impacts on water demand, with further impact on the ecosystems. Under all climate
change scenarios in the Mediterranean region, available water resources decrease
while irrigation demand increases (Iglesias et al. 2007).
Climate change drives much of the change evident in natural hydrological cycles,
which is one of the greatest environmental, social and economic threats facing the
planet. Recent warming of the climate system, irrespective of the causes is indisput-
able, and is now evident from observations of increases in global average air and
ocean temperatures, widespread melting of snow and ice, and rising global mean
sea level. IPCC (2007) concludes that observational evidence from all continents
and most oceans shows that many natural systems are being affected by regional
climate changes, particularly temperature increases. Other effects of regional climate
changes on natural and human environments are emerging, although many are
difficult to discern due to adaptation and non-climatic drivers.
Anticipated impacts of climate change on fresh water resources and their man-
agement are reported to be as follows (IPCC 2007):
• By mid-century, annual average river runoff and water availability are projected
to increase by 10–40% at high latitudes and in some wet tropical areas, and
decrease by 10–30% over some dry regions at mid-latitudes and in the dry trop-
ics, some of which are presently water stressed areas. In some places and in
particular seasons, changes differ from these annual figures.
• Drought-affected or water stressed areas will likely increase in extent.
• Heavy precipitation events are very likely to increase in frequency and intensity,
and thus to augment flood risks.
• In the course of the century, water supplies stored in glaciers and snow cover are
projected to decline, reducing water availability in regions supplied by meltwater
from major mountain ranges, where more than one-sixth of the world population
currently lives.
Impacts of climate change are of diverse nature. They refer to losses in biodiver-
sity due to changes in environmental conditions affecting the ecosystems. The pres-
ent boundaries of natural ecosystems may change due to modifications in climate
regimes; actual crop patterns may have to be modified due to changes in environ-
mental conditions influencing the crop cycles, development and production. Rainfed
crops are therefore more vulnerable than irrigated ones due to changes in precipita-
tion, infiltration, evapotranspiration and soil moisture regimes. Food security is
therefore threatened in more vulnerable regions and countries of the world.
Changes in rainfall regimes will induce changes in stream flow regimes and lower
base flow is expected. Moreover, the water quality regimes will also change and
contamination impacts may be larger, affecting public health. The latter may also be
impacted due to increase of frequency and severity of heat waves and wildfires.
Overall, the water availability is expected to decrease thus enhancing competition
among users and making it more difficult to satisfy the increased urban water
demand for residents and tourism. It is important to recognize climate change as a
process driving exacerbated water scarcity and threatening development in develop-
ing countries. Unfortunately, many other processes and driving forces are contribut-
ing to degradation of Earth’s environment and people’s welfare, including devastating
wars. Nowadays, it is possible to identify within regions several situations that are
expected to arise due to climate change (Pereira et al. 2009).
Water scarce regions are highly vulnerable to climate change impacts. Coping
with water scarcity involves therefore requires that such impacts be recognized and
appropriate mitigation and adaptation measures be developed and implemented to
effectively cope with it. In general with the increase in temperature it is likely that
there will be an increase of potential evapotranspiration and therefore a higher veg-
etation and crops demand for water as well as impacts due to heat waves.
Briefly, the impacts of climate change on fresh water resources and their man-
agement can be concluded that the annual average river runoff and water availability
are projected to increase by 10–40% at high latitudes and in some wet tropical areas,
and decrease by 10–30% over some dry regions at mid-latitudes and in the dry
tropics. Drought-affected or water stressed areas will likely increase in extent and
heavy precipitation events are very likely to increase in frequency and intensity,
and thus to augment flood risks. And finally, water supplies stored in glaciers and
snow cover are projected to decline, reducing water availability in regions supplied
by meltwater from major mountain ranges, where more than one-sixth of the
world population currently lives.
4.5 The Challenges Egypt Faces with Regard to Water

and Agricultural Development
Egypt, like any other arid country, faces the pressing challenge of closing the gap
between its limited water resources and the increasing water demand. Egypt consid-
ers the River Nile its ‘vein of life’, being the sole source that covers nearly 95% of
the population requirements. The dependence of the other nine riparian countries on
the Nile water varies according to each country’s precipitation and water use pat-
terns. These countries, being mostly humid and/or less populated than Egypt, are
less vulnerable to fluctuations of the Nile flows. Despite the fact that declining Nile
water availability with respect to growing populations and increasing requirements
for development is an alarming issue, Egypt has not yet reached the stage of a
crisis. The principal water management challenges in Egypt stem from the nature
and quality of supply and demand management responses to water shortage.
Table 4 Estimated water balance (in km3 year−1) of Egypt in 1997 and 2017 from Ministry of
Water Resources and Irrigation through making a comparison between resources and demand
water (Adapted from Adly and Ahmed 2009)
Water resources Water demand
Items 1997 2017 Items 1997 2017
From lake Nasser through high 55.5 55.5 Agriculture 57.8 63.6
Aswan dam
Rainfall 1.3 1.3 Domestic 4.7 6.6
Shallow groundwater 6.1 8.4 Industry 7.5 18.7
Drainage reuse 7.5 11.4 Navigation 0.2 0.2
Wastewater reuse 1.4 2.4 Evaporation 2.4 2.5
Total 71.8 79.0 Fishery 1.3 0.6
Industrial water flushed back to system 6.8 17.8 Total 73.9 92.2
Agricultural water flushed back to 4.9 1.9 Drainage to sea 12.9 9.5
system (not including reuse)
Domestic water flushed back to system 2.4 2.6
(not including reuse)
Fishery water flushed back to system 0.9 0.4
Total water resources 86.8 101.7 Total water 86.8 101.7
demand
Table 4 shows the water demand in Egypt in 1997 and that projected for 2017, dem-
onstrating how these requirements will be met through tapping non-conventional
water resources, including water savings and possibilities of reuse. Water conditions
are, however, likely to straighten despite the giant water storage reservoir in Lake
Nasser created after the construction of the High Aswan Dam in 1964. This is due
to climatic fluctuations, accelerating development activities, and high price of untra-
ditional water abstractions. The role of the High Aswan Dam in reducing Egypt’s
vulnerability to the fluctuations of external and shared resources originating from
upstream countries cannot, nonetheless, be denied (Adly and Ahmed 2009).
The challenges facing all stakeholders led to recommendations based on mani-
fold expertise and a willingness to start where others have ended. The cooperation
within the Nile Basin Initiative offers a platform for partnership and a shared vision.
There is a need to develop national integrated programs for water resources man-
agement. The civil society has an important role to play especially in the areas of
water conservation and sustainable livelihoods through food security. And it would
be appropriate in this context to recall the goal of the Nile Basin Discourse “to pro-
mote dialogue on sustainable and equitable development, peace and mutual under-
standing within the Nile River Basin”. These data point out the need for coordination
and effort unification among all Nile basin countries to reach unconventional solu-
tions in facing the problem of the declining individual share of water supply, includ-
ing improved water conservation, waste minimization and reuse of treated sewage
water, besides a rationalization of the water demand (Adly and Ahmed 2009).
Managing water resources will become a more complex endeavor with climate
change. Analysis predicts that climate change will intensify and accelerate the
hydrological cycle, which will result in more water being available in some parts of
the world and less water being available in other parts of the world (most of the
developing world). Weather patterns are predicted to be more extreme. Those
regions adversely affected will experience droughts and/or possible flooding. Is
Egypt vulnerable? The answer is yes. The Nile waters are highly sensitive to climate
change, both in amount of rainfall and variations in temperature. And since these
two factors are also interrelated, i.e., temperature changes affecting rainfall, it can
be expected that climate change will take the form of changes in levels of precipita-
tion as a result of changes in temperature, or other factors, and that the resulting
effect on the Nile flows will be from moderate to extreme, with the latter scenario
most likely in the long term (Elsaeed 2012).
In conclusion, Egypt faces the pressing challenge of closing the gap between its
limited water resources and the increasing water demand. Egypt considers the River
Nile its vein of life, being the sole source that covers nearly 95% of the population
requirements. The Nile waters are highly sensitive to climate change, both in amount
of rainfall and variations in temperature. And since these two factors are also inter-
related, i.e., temperature changes affecting rainfall, it can be expected that climate
change will take the form of changes in levels of precipitation as a result of changes
in temperature, or other factors, and that the resulting effect on the Nile flows will
be from moderate to extreme, with the latter scenario most likely in the long term.
4.6 Climate Changes and Nile Water Availability in Egypt
Any assessment of Egypt’s water resources recognizes the country’s enormous reli-
ance on the Nile, which makes up about 95% of Egypt’s water budget. Other sources
of Egypt’s water budget, precipitation and ground water, do not make up more than
5% of the available supply, although the effect of increases or decreases in precipita-
tion near the sources of the Nile can have a larger than expected effect on Nile flows.
Egypt’s total water budget is produced by a combination of three variables: the Nile
(95%), precipitation (3.5%) and ground water (1.5%). The Nile produces 55.5 bil-
lion m3, while the latter two variables combine to form safely about 2.2 billion m3
of fresh water. In total, Egypt has available fresh water reserves of 58 billion m3.
Egypt’s annual water demand is about 77 billion m3. The deficit between Egypt’s
water supply and demand must be met through recycling. The 19 billion m3 deficit
is filled by a combination of treated sewage and industrial effluent, i.e. 4 billion m3
and recycling used water, mainly from agriculture, i.e. 8 billion m3. An additional
4 billion m3 is extracted from the shallow aquifer and 3 billion m3 comes from the
Al Salam Canal Project (Elsaeed 2012).
Recycling is partly natural and partly intentional. Water reclaimed from agricul-
ture is a natural process of drainage waters returning to the Nile. The remaining
two sources of recycled water, the Al Salam Canal and extraction from the shallow
aquifer, are manmade solutions to the deficit. Consumption of the 77 billion m3 in

annual water demand in Egypt is mainly from agriculture, i.e. 62 billion m3. An
additional 10%, i.e. 8 billion m3 is used as drinking water. Approximately 95% of
the population relies on this water for drinking purposes. The remaining demand
comes from industry, i.e. 7.5 billion m3. This section will focus on the impact of
climate change on water supply and the potential challenges Egypt will face in the
future if the balance between water supply and demand is altered (Elsaeed 2012).
Egypt’s climate is semi-desert, characterized by hot dry summers, moderate winters,
and very little rainfall. The country has areas with strong wind, especially along the
Red Sea and Mediterranean coasts. Sites with an annual average wind speed of 8.0–
10.0 m sec−1 have been identified along the Red Sea coast and about 6.0–6.5 m sec−1
along the Mediterranean coast. Average precipitation in the Ethiopian highlands,
where much of the water in the Nile originates is highest in July, August, and
September, at 5.4 mm day−1, and almost negligible between January and March.
Egypt is fairly unique in the distribution of its population, land-use and agriculture,
and economic activity which makes it extremely vulnerable to any potential impacts
on its water resources and coastal zone. Despite being a large rectangular shaped
country with an area of about a million square kilometers, its lifelines are con-
strained along a narrow T-shaped strip of land, which constitutes less than 5% of its
land area along the Nile and the coast around the Nile delta. The Nile supplies 95%
of the country’s total water needs, including water intensive irrigated agriculture
along its banks and the delta. Agriculture is quite critical to the national economy as
it employs 30% of the work force and contributes 17% to the Gross National Product
(GNP). Major urban centers, commerce, and industrial activity are also confined to
the narrow corridor along the Nile and the coast around its delta. The rest of the
country is desert and does not support much population or economic (Agrawala
et al. 2004)
The potential impacts of climate change on coastal resources are ranked as most
serious. Sea levels are already rising in the Nile delta due to a combination of factors
including coastal subduction and reduced sediment loads due to the construction of
the High Aswan Dam upstream. Climate change induced sea-level rise only rein-
forces this trend. In addition to this high biophysical exposure to the risk of sea level
rise, Egypt’s social sensitivity to sea level rise is particularly high. As discussed
earlier in this section much of Egypt’s infrastructure and development is along the
low coastal lands and the fertile Nile delta also constitutes the prime agricultural
land in Egypt. The loss of this land due to coastal inundation or to saline intrusion
will therefore have a direct impact on agriculture, which in turn is critical to Egypt’s
economy.
Egypt’s Nile delta with its coastal front on the Mediterranean is considered vul-
nerable to the impacts of climate change. In addition to expected rise in sea-level,
shoreline erosion, stresses on fisheries and saltwater intrusion in groundwater create
major challenges. These factors also produce stressful effects on water and agricul-
tural resources, tourism and human settlements. Fragile and unique ecosystems
such as the mangrove stands in the Red Sea, which stabilize shorelines and provide
a habitat for many species, may also be threatened. The northern Egyptian lakes,
which constitute about 25% of the total Mediterranean wet lands and produce about
60% of the fish products, are also highly vulnerable to the impacts of climate change.
Since the lakes are relatively shallow, climate change can lead to an increase in
water temperature, which could result in changes in the lake ecosystems as well as
changes in yield. So far, in-depth studies on potential impacts of climate change on
lake ecosystems are not available (Agrawala et al. 2004).
It cold be summarized that Egypt’s total water budget is produced by a combina-
tion of three variables: the Nile (95%), precipitation (3.5%) and ground water
(1.5%). The Nile produces 55.5 billion m3, while the latter two variables combine to
form safely about 2.2 billion m3 of fresh water. In total, Egypt has available fresh
water reserves of 58 billion m3. Egypt’s annual water demand is about 77 billion m3.
The deficit between Egypt’s water supply and demand must be met through recy-
cling. The 19 billion m3 deficit is filled by a combination of treated sewage and
industrial effluent, i.e. 4 billion m3 and recycling used water, mainly from agricul-
ture, i.e. 8 billion m3. An additional 4 billion m3 is extracted from the shallow aqui-
fer and 3 billion m3 comes from the Al Salam Canal Project.
4.7 The Impacts of Sea Level Rise on Egypt
Vulnerability to climate change is considered to be high in developing countries due

to social, economic, and environmental conditions that amplify susceptibility to
negative impacts and contribute to low capacity to cope with and adapt to climate
hazards. Moreover, projected impacts of climate change generally are more adverse
for lower latitudes, where most developing countries are located, than for higher
latitudes. Because of the high level of vulnerability, there is an urgent need in the
developing world to understand the threats from climate change, formulate policies
that will lessen the risks and to take action. The danger is greatest, where natural
systems are severely degraded and human systems are failing and therefore inca-
pable of effective response, specifically in deprived nations. Moreover, land degra-
dation and desertification may also be exacerbated in these areas, posing additional
threats to human well-being and development, added by intensified human pres-
sures on lands and poor management. The livelihoods and food security of the rural
poor are threatened by climate change with all its impacts, and the vulnerability to
adverse health impacts is greater where health care systems are weak and programs
for disease surveillance and prevention are lacking. In addition to multiple factors
converging to make the people inhabiting coastal zones and small islands highly
endangered from the causes of sea level rise. Egypt’s coastal zone of the Nile delta
has been defined as a vulnerable zone as a consequence of sea level rise combined
with geological and human factors (El-Sharkawy et al. 2009)
It is well documented that sea level changes are caused by several natural phe-
nomenon; the three primary contributing ones are: ocean thermal expansion, glacial
melt from Greenland and Antarctica -in addition to a smaller contribution from
other ice sheets- and change in terrestrial storage. Among those, ocean thermal
expansion has been expected to be the dominating factor behind the rise in sea level.
However, new data on rates of deglaciation in Greenland and Antarctica suggest
greater significance for glacial melt, and a possible revision of the upperbound
estimate for sea level rise in this century. It is predicted that, with global warming,
global average sea levels may rise by between 7 and 36 cm by the 2050s, by between
9 and 69 cm by the 2080s and 30–80 cm by 2100. The majority of this change will
occur due to the expansion of the warmer ocean water. Since the Greenland and
Antarctic ice sheets contain enough water to raise the sea level by almost 70 m,
people will be directly affected by rising sea levels in several ways. As seas rise
many areas of the coasts will be submerged, with increasingly severe and frequent
storms and wave damage, shoreline retreat will be accelerated. In addition to
expected disastrous flooding events caused by severe climate events such as heavy
flooding, high tides, windstorms in combination with higher seas (Dasgupta et al.
2007). The impacts of sea level rise will not be globally uniform, because of local
variations in vertical crustal movements, topography, wave climatology, long shore
currents, and storm frequencies. Low gradient coastal landforms most susceptible to
inundation include deltas, estuaries, beaches and barrier islands, and coral reefs.
Regions at risk include the Low Countries of Europe, eastern England, the Nile
delta in Egypt, the Ganges–Brahmaputra, Irrawaddy, and Chao Phraya deltas of
south-eastern Asia, eastern Sumatra, and Borneo. In the United States, the mid-
Atlantic coastal plain, the Florida Everglades, and the Mississippi delta will be par-
ticularly vulnerable (Vivian 2005)
The Nile Delta is one of the oldest intensely cultivated areas on earth. It is very
heavily populated, with population densities up to 1,600 inhabitants per square kilo-
meter. The low lying, fertile floodplains are surrounded by deserts. Only 2.5% of
Egypt’s land area, the Nile delta and the Nile valley, is suitable for intensive agricul-
ture. Most of a 50 km wide land strip along the coast is less than 2 m above sea-level
and is protected from flooding by a 1–10 km wide coastal sand belt only, shaped by
discharge of the Rosetta and Damietta branches of the Nile. Erosion of the protec-
tive sand belt is a serious problem and has accelerated since the construction of the
Aswan dam (Fig. 5). Rising sea level would destroy weak parts of the sand belt,
which is essential for the protection of lagoons and the low-lying reclaimed lands.
The impacts would be very serious: One third of Egypt’s fish catches are made in
the lagoons. Sea level rise would change the water quality and affect most fresh
water fish. Valuable agricultural land would be inundated. Vital, low-lying installa-
tions in Alexandria and Port Said would be threatened. Recreational tourism beach
facilities would be endangered and essential groundwater would be salinated. Dykes
and protective measurements would probably prevent the worst flooding up to a
50 cm sea level rise. However, it would cause serious groundwater salination and
the impact of increasing wave action would be serious.
The Nile Delta is 200 km long and 255 km wide, within a coastline of over
1,000 km on the Mediterranean Sea. The low sandy coast of the Nile Delta stretches
with an arc between Ras Abu Quir to the west and the Bay of Tinah, to the east.
Fig. 5 Nile Delta: Potential impact of sea level rise in 2002 and the situation for the Nile Delta
when this sea level rise reaches to 0.5 m and 1.0 m. That means at 0.5 m and 1.0 m, the damage for
cropland will be 1,800 and 4,500 km2, respectively. At the same time the population will be dis-
placed at rate of 3.8 and 6.1 million inhabitant, respectively. Source: Simonett (2002), in: UNEP/
GRID-Arendal Maps and Graphics Library (Retrieved 15:46, January 13, 2012 from http://maps.
grida.no/go/graphic/nile_delta_potential_impact_of_sea_level_rise)
Two branches of the Nile have formed the promontories at Rosetta and Damietta.
Egypt’s second largest city, Alexandria is located on the northwestern part of the
coastal delta zone, with a population of 3.3 million in 1996, and more than 4.1 mil-
lion in 2006 (CAPMAS 2006 census).
Alexandria is the main harbor of Egypt and hosts around 40% of the country’s
industrial capacity, in addition to being an important summer resort and trading
centre. Other large cities in the northern, low-lying delta zone include the rapidly
growing city of Damietta and the historic city of Rosetta and Port Said City to the
eastern side of the delta. The Nile delta region is fairly unique in the distribution of
its population, topography, land-use, agricultural productivity and economic activi-
ties, which makes it extremely vulnerable to any potential impacts on its water
resources and coastal zone (El Raey 2011).
The River Nile supplies 95% of the country’s total water needs, including water
intensive irrigated agricultural land along its banks and the delta. Agriculture is
quite critical to the national economy as it employs 30% of the work force and con-
tributes 17% to the gross domestic production (IDSC 2009). Major urban centers,
commerce, and industrial activities are also confined to the narrow corridor along
the Nile and the coast around its delta. The rest of the country (95%) is desert and
does not support much population or economic activity. The Nile Delta region lies
within the temperature zone, which is a part of the great desert belt. The average
temperatures in January and July in Cairo are 12°C and 31°C, respectively. Minimum
and maximum temperatures in Cairo are 3°C and 48°C, respectively. Rainfall over
the Nile Delta is rare and occurs in winter. Maximum average rainfall along the
Mediterranean Sea shore, where most of the rain occurs, is about 180 mm. This
amount decreases very The Nile delta region is the most fertile land of Egypt which
depends mainly on water that reaches the region through the River Nile with
resources on the Ethiopian hills and Lake Victoria some several thousand kilome-
tres to the south. The Nile delta coast stretches about 300 km and hosts a number of
highly populated deltaic cities such as Alexandria, Port-Said, Rosetta, and Damietta.
These cities are also critical centers of industrial and economic activity. In addition,
the Nile delta coastal zone includes a large portion of the most fertile low land of
Egypt. The topography is generally sloping from the apex at Cairo to the
Mediterranean coast at a rate of about 1 m km−1 with varying sand dunes, ridges and
low elevation areas near the coast. The coastal zone of Egypt hosts five northern
lakes which constitute about 25% of the wetland of the Mediterranean and are con-
sidered main sanctuaries for birds and fish resources (El Raey 2011).
There are conflicting projections of the future availability of the water of the Nile
as a result of climate change. Yates and Strzepek (1998), using a monthly water bal-
ance model, reported that five of six global circulation models (GCMs) showed for
doubled CO2 levels increased flows at Aswan, with increases of as much as 137%
(United Kingdom Meteorological Organization). Only one global circulation model
(GFDLT) showed a decline in annual discharge at Aswan (−15%). The variations of
the results indicate that more robust studies are needed to provide a more solid base
for the design of public policy. However, the more plausible projections seem to
point to a reduced availability of Nile water for Egypt in the future. In addition,
El Shamy et al. (2009) confirmed this strong uncertainty using 17 IPCC models.
This global sea-level rise combined with local land subsidence in many coastal
areas, are expected to cause serious damage to many coastal ecosystems espe-
cially those of the low land deltaic coasts such as that of the Nile Delta in Egypt
(El Raey 2011).
It could be concluded that vulnerability to climate change is considered to be
high in developing countries due to social, economic, and environmental conditions
that amplify susceptibility to negative impacts and contribute to low capacity to
cope with and adapt to climate hazards. The sea level changes are caused by several
natural phenomenon; the three primary contributing ones are: ocean thermal expan-
sion, glacial melt from Greenland and Antarctica -in addition to a smaller contribu-
tion from other ice sheets- and change in terrestrial storage. Rising sea level would
destroy weak parts of the sand belt, which is essential for the protection of lagoons
and the low-lying reclaimed lands. The impacts would be very serious: One third of
Egypt’s fish catches are made in the lagoons. Sea level rise would change the water
quality and affect most fresh water fish.
4.8 Impact of Climate Change on Crop Production in Egypt
Globally, agricultural emissions have increased by 14% from 1990 to 2005 with an
average annual emission of 49 Mt CO2 eq. yr−1 (US-EPA 2006). N2O from soils and
manure management and CH4 from enteric fermentation were the agricultural
sources, showing the highest increase in emissions at 21%, 18% and 12%, respec-
tively. N2O emissions increased by 31 Mt CO2 yr−1, which is almost twice the rate of
increase for CH4 emissions. United State Environmental Protection Agency fore-
casts acceleration in the global greenhouse gases emission from agriculture for the
period 2005–2020. In the developing countries, the growth is expected to continue
at the same rate as in 1990–2005, whereas in the more developed regions, the
decreasing trend would be reversed and emission would grow by 8% up to 2020
(US-EPA 2006). Two most significant sources, N2O from soils and CH4 from enteric
fermentation, would also increase quite rapidly. N2O emission, which is expected to
an average of 49 Mt CO2 yr−1, would continue to grow faster than CH4 emissions,
projected to an average of 35 Mt CO2 yr−1 (Adhya et al. 2009).
Specific management options can be used to reduce agriculture’s environmental
impacts. Conservation practices, that help prevent soil erosion, may also sequester
soil C and enhance CH4 consumption. Managing N to match crop demands can
reduce N2O emission, while manipulating animal diet and manure management can
reduce both CH4 and N2O emission from animal husbandry. Thus, all segments of
agriculture have the management options which can reduce agriculture’s greenhouse
gases footprints. Opportunities for mitigating greenhouse gases emissions in agricul-
ture can be grouped into three broad categories based on the following principles:
• Reducing emissions: The fluxes of greenhouse gases emissions can be reduced
by managing more efficiently the flows of carbon and nitrogen in agricultural
systems. The exact approaches, that best reduce emissions, depend on local
conditions and therefore, vary from region to region.
• Enhancing removals: Agricultural ecosystems hold large reserves of C, mostly in
soil organic matter. Any practice, that increases the photosynthetic input of C or
slows the return of stored C via respiration, will increase stored C, thereby
‘sequestering’ C or building C ‘sinks’.
• Avoiding emissions: Using bioenergy feed-stocks would release CO2-C of recent
origin and would, thus, avoid release of ancient C through combustion of fossil
fuels. Emissions of greenhouse gases emissions can also be avoided by agricul-
tural management practices that forestall the cultivation of new lands (Adhya
et al. 2009).
Agriculture in Egypt is expected to be especially vulnerable because of hot cli-
mate. Further warming is consequently expected to reduce crop productivity. These
effects are exacerbated by the fact that agriculture and agro-ecological systems are
especially prominent in the economics of Egypt as one of the African countries. The
rapid growth of the country’s population, the economic stress of reliance on food
imports, and the limited area for agriculture requires finding new ways to increase
agricultural productivity in general and oil crops in specific. If climate change as
projected by atmospheric scientists adversely affected crop production, Egypt would
have to increase its reliance on costly food imports.
The potential impact of climate change on some field crops production and
evapotranspiration in Egypt was studied through DSSAT3 and DSSAT3.5 (Tsuji
et al. 1995, 1998), and COTTAM (Jackson et al. 1988) models, (El-Shaer et al.
1997; El-Marsafawy et al. 2007; El-Marsafawy 2007). Based on the mentioned pre-
vious simulation studies, climate change could decrease national production of
many crops (ranging from −11% for rice to −28% for soybean) by the year of 2050
compared to their production under current conditions. Yield of cotton would be
increased in comparison with current climate conditions. At the same time, water
consumptive use for summer crops will be increased up to 8% for maize and up to
16% for rice by the year 2050 compared to their current water consumption
(El-Marsafawy and El- Samanody 2009).
To investigate the impacts of climate change on sunflower productivity, water
consumptive use, crop water productivity, farm net return and how to mitigate the
potential effects of climate change on this crop, El-Marsafawy and El- Samanody
(2009) studied the economic impacts of future climatic changes on sunflower crop
in Egypt. They concluded that, climate change could decrease sunflower seed yield
by 27%, increase water consumptive use by 12% and decrease crop water produc-
tivity accordingly by 34%. Changing sowing date of sunflower from 1st to 10th of
June to 1st to 10th of May could increase seed productivity about 13–18%. Reducing
irrigation water amounts by 10% could be recommended as a way to conserve irri-
gation water without clear reduction in seed yield. Climate change without adapta-
tion studies could decrease farm net return about 44 and 63% for holders who own
the land, and holders who rent it, respectively. At the same time, climate change
could decrease the economic return from the water unit about 35%.
In brief, opportunities for mitigating greenhouse gases emissions in agriculture

can be grouped into three broad categories based on reducing emissions, enhancing
removals and avoiding emissions. Agriculture in Egypt is expected to be especially
vulnerable because of hot climate. Further warming is consequently expected to
reduce crop productivity. These effects are exacerbated by the fact that agriculture
and agro-ecological systems are especially prominent in the economics of Egypt as
one of the African countries. In Egypt, climate change could decrease sunflower
seed yield by 27%, increase water consumptive use by 12% and decrease crop water
productivity accordingly by 34%.
5 Conclusion
In the beginning of the current century, the world is facing critical global food and
fuels shortages, climate change, urban growth, environmental degradation, and nat-
ural disaster-related challenges as today’s world population continues to grow.
Today the entire world is aware that our food supply cannot meet the demands of the
world population. How can that be possible? Why can’t we feed the people of the
world? The answer lies in agricultural sustainable development. Agriculture is a
major economic issue in Egypt. It is an issue as a local food source, for international
trade, for balance of payments, land use and water use and as a basic product for
food and fiber manufacturing. Hence, every aspect of the economic structure of the
country relates to agriculture. For over 5000 years the farmers of Egypt created a
civilization based on the union of the land and the Nile River. It was one of the earli-
est civilizations and it had a profound influence on the region. Agriculture created
most of Egypt’s wealth. Egypt, because of its very limited arable land and water
resources, is probably more dependent on research to expand food production than
any other country in the world. In Egypt, the search for ways to achieve sustainable
agriculture and natural resource management requires changes in the traditional
approach to problem solving. Researchers must cross the boundaries of their indi-
vidual disciplines; they must broaden their perspective to see the merits of indige-
nous knowledge; and they must look to the farmer for help in defining a practical
context for research. This change in vision is under way in various degrees through-
out the research community, but the pace of change is slow. Two key indicators of
deterioration in agricultural systems are declines in the quality of the soil and of the
water. Poor management of either of these resources quickly leads to decreases in
farm productivity. However the lack of any emphasis on extension of the resulting
information continues to be the major problem for Egypt and the region associated
with the use of the Nile waters. Egypt is subject to potential impacts of climate
change, including sea level rise, inundation of the low lying lands in the Nile Delta
that could reach 10–12% of the total area, impacts on water resources and agricul-
tural productivity and associated social and economic effects. If Egypt’s continued
high rate of natural population increase is not reduced it could result in a population
of 140 million inhabitants by 2050. This challenge calls for the need to develop a
new vision for Egypt that mainly aims to deal with the potential link between climate
change (including SLR, water scarcity and desertification) and human mobility
(including displacement) in Egypt.
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Adapting Maize Crop to Diverse
Agro-Ecosystems
Ioannis S. Tokatlidis
Abstract Global climate change compels agriculture to be productive under diverse

and marginal conditions. In maize, modern hybrids fail to meet this requirement.
Although breeding has achieved spectacular progress in grain yield per area through
improved tolerance to stresses including intense crowding, yields at low plant popu-
lation densities remain almost unchanged. Stagnated plant yield potential renders
hybrids unable to take advantage of resource abundance at lower populations, des-
ignating them population-dependent. Consequently, the optimum population varies
greatly across environments. Generally, the due population increases as the environ-
mental yield potential gets higher. As a remedy, relatively low populations are
recommended for low-input conditions leading to inappropriate population in occa-
sional adequacy of resources and considerable yield loss. For example, for a rain-fed
hybrid tested at one location across 11 seasons, crop yield potential and optimum
population on the basis of the quadratic yield-plateau model varied from 1,890–
8,980 kg/ha and 4.56–10.2 plants/m2, respectively; grain yield loss of 1,600 kg/ha is
computed for a favorable season if the lowest optimum population is used.
This article reviews the consequences in terms of crop sustainability under
widely diverse environments imposed by climatic changes. This article then pro-
poses crop management strategies to address the situation. The major points are: (1)
variable-yielding environments require variable optimum populations, (2) population
dependence is an insurmountable barrier in making a decision on plant population,
(3) farmers suffer from considerable yield and income loss, (4) estimating the less
population-dependent hybrids among the currently cultivated ones is a major chal-
lenge for agronomists, and (5) the development of population-neutral hybrids is a
fundamental challenge for maize breeding. The honeycomb breeding is a valuable
tool to pursue this goal, since it places particular emphasis on the so-far stagnated
plant yield potential that is essential for population-neutral hybrid development.
I.S. Tokatlidis (*)

Department of Agricultural Development, Democritus University of Thrace,
Pantazidou 193, Orestiada 68200, Greece
e-mail: itokatl@agro.duth.gr; itokatl@hotmail.com

98 I.S. Tokatlidis
Keywords Crop yield potential • Honeycomb breeding • Optimum population

• Plant yield potential • Population-neutral hybrids • Sustainable agriculture
1 Introduction
Tremendously variable weather conditions arise from climatic changes, and the
pace of future environmental change will likely be unprecedented (Cutforth et al.
2007). It is expected that these weather events will have implications for agro-
ecosystems, with crop yields becoming more variable (Lavalle et al. 2009). Consequently,
agriculture to be sustainable in the future must be able to be adequately productive
under diverse and marginal conditions. Moreover, agricultural systems are under
increasing pressure to supply food to a growing human population (Hatfield et al.
2011; Jégo et al. 2011), and food demand globally is expected to double by 2,050
(Stuber and Hancock 2008). Flexibility of agriculture has been highlighted as a
determinant factor of sustainability, enabling agriculture to adapt to ongoing envi-
ronmental changes and allowing the preservation of the ability to farm and produce
food into the future (Lichtfouse et al. 2009). However, there are serious concerns
that the forthcoming environmental changes will make the challenge of feeding
additional people exceedingly difficult within the next 50 years (Vadez et al. 2012).
In sequence, new cultivars, cropping systems, and agricultural management strate-
gies are needed to provide options to farmers to counterweigh these changes.
At present, maize is of the highest tonnage cereal crops worldwide, providing
feed, food, and fuel for more than 6,000 million humans, while unprecedented growth
in global demand for cereals is expected (Troyer and Wellin 2009). However, the
crop suffers from an agronomic weakness of prime significance, affecting its grain
productivity and stability. Modern hybrids (Fig. 1) are usually population-dependent
(Tokatlidis et al. 2001, 2011), with the ideal plant number per area depending on
several factors, including water availability, soil fertility, hybrid maturity group, and
row spacing (Sangoi et al. 2002). Yet hybrids accomplish their per area yield poten-
tial at high and narrow spectrum of populations, i.e. they follow the quadratic-plateau
regression model (Van Roekel and Couter 2011). Tokatlidis and Koutroubas (2004)
reviewed the adverse effects of indispensable high populations on grain yield stabil-
ity because of considerable yield loss due primarily to missing plants, increased
plant-to-plant variability, raised stalk lodging, and augmented barrenness.
This chapter is an abridged version of a review article entitled “Adapting maize
crop to climate change” (Tokatlidis 2013). It deals with the issue of great variability
in optimum populations as a consequence of the strong hybrid by population interac-
tion. The main hypothesis comprises: (1) yield potential varies across locations and/or
seasons, and the same applies for optimum population, (2) hybrids usually fail to
meet the requirements of the diversified environments due to their capacity to attain
yield potential at a particular population, resulting in yielding penalty, (3) hybrids
that accomplish their crop yield plateau at a relatively wide range of populations are
more flexible, and (4) low threshold of a wide spectrum of optimum population is a
determinant of ideal hybrids for flexible agriculture under variable conditions.
Adapting Maize Crop to Diverse Agro-Ecosystems 99
Fig. 1 Modern maize hybrids usually accomplish their per area yield potential, i.e. crop yield
potential, at high and narrow spectrum of populations with optimum population depending on
climate and availability of resources, thereby designating them population-dependent
Data were obtained from a number of the most recent papers dealing with maize
grain yield response to population, particularly those including variable environments
and different hybrids. Key measures, either provided or estimated from the available
data, were: (1) the environmental/hybrid grain yield potential at the area level through
either the experimental mean yield or the max attainable yield, i.e. crop yield potential,
(2) the optimum plant population density to effectively exploit resources at the per-area
level, and (3) the grain yield potential at the single-plant level, i.e. plant yield potential.
The per unit ground area maximum yield and the required number of plants were com-
puted from the quadratic equation of yield response to plant population, corresponding
to the crop yield potential and optimum population, respectively. Plant yield potential,
defined as the maximum yield per plant when any kind of stress is absent, i.e. in the
absence of competition, was measured indirectly by the Yan and Wallace (1995) proce-
dure, i.e. through the intercept of the linear regression analysis of yield per plant over
plant population (Tokatlidis 2001; Tokatlidis and Tsialtas 2008).
2 Interactions Among Environments, Hybrids, and Populations
From studying research works dealing with different hybrids it becomes apparent
that hybrids may respond differently to plant population changes. Moreover, the
higher the yield potential of the environment, the higher the due population should
100 I.S. Tokatlidis
be. Simply, the optimum population may differ not only among different hybrids,
but even for the same hybrid across environments, i.e. across locations or across
seasons at the same location. Thus, it is hard to reach a particular recommendation
on appropriate plant population, devoid of the risk of considerable yield loss and
limitation of farmers’ income.
2.1 Implications of Environmental Variability

on Optimum Population
The problem of population dependence becomes more acute when the crop is produced
under rain-fed conditions. Dryland maize production is usually carried out at diverse
conditions, reflecting an analogous diversity in optimum population. The interaction
among hybrid, population, and environment constitutes the root cause of the variability
in optimum population among different hybrids and, more importantly, of the wide vari-
ability in optimum population for the same hybrid across seasons and/or locations.
Blumenthal et al. (2003) reported the results of a 2-year study (1999, 2000) aim-
ing to determine appropriate population for dryland maize grown in western
Nebraska, USA. Field studies were conducted across four locations, where the
hybrid PR3893 was no-till seeded into wheat or proso millet stubble at the popula-
tion range of 1.73–5.68 plants/m2. The experimental mean yield ranged from 1,220
to 5,550 kg/ha, implying more than 300% yield gap across locations and seasons.
The respective quadratics gave a tremendous range of optimum population (i.e.
0.14–10.3 plants/m2). These calculations might be exaggerated and beyond the
expected limits. Nevertheless, they pinpoint the question of variation in optimum
population on account of the environmental variance in yield potential.
Across 11 consecutive growing seasons (from 1989 to 1999) at the same location
(Martonvásár, Hungary), rain-fed maize crop averaged a grain yield from 1,460 to
7,670 kg/ha over four N-fertilizer treatments at 3, 5, 7, and 9 plants/m2, showing an
experimental mean yield gap of up to 425% across seasons (Berzsenyi and Tokatlidis
2012). The corresponding range of crop yield potential was 1,600–9,130 kg/ha, and
concerning the optimum population the ideal plant number per square meter
was 5.29 for the driest 1990 season and within the 5.33–13.2 range for the remaining
10 seasons (Fig. 2).
The phenomenon of variation in optimum population is presumably more inten-
sive in widely diverse environments. Nevertheless, substantial variation in optimum
population is apparent even from studies conducted in less diverse environments.
Farnham (2001) evaluated the hybrid N4640Bt at six locations in Iowa, USA. Over
four plant populations, two row spacing, and three seasons, the across location experi-
mental mean yields were from 9,950 up to 11,060 kg/ha, implying a gap in
experimental mean yield of only 11%. However, a high range of optimum population
values was derived from the quadratic regression analysis, i.e. 7.6–14 plant/m2.
With the main objective to determine the optimum population for maize in
Wisconsin, USA, Stanger and Lauer (2006) evaluated hybrids with non-Bt and
Fig. 2 Variability in crop yield potential is indicative of the across-season at a single-location

environmental diversity in dryland maize production, accompanied by analogous variation in
optimum population. The linear correlation between the two measures is positive (r = 0.67,
P < 0.03). Data over four hybrids and four N-fertilizer treatments at four populations from
Berzsenyi and Tokatlidis (2012)
Bt traits at the target populations of 6.18–12.35 plants/m2 for three growing seasons
(2002–2004) and across 10 locations belonging to three maize zones. They pre-
dicted for the non-Bt hybrid group in particular an across-location crop yield
potential varying from 8,900 to 15,200 kg/ha, accompanied by a variation in opti-
mum population from 5.78 to 11.5 plants/m2.
Popp et al. (2006) evaluated the profit-maximizing plant population of several
maize hybrids at two locations of Arkansas, USA, i.e. Fayetteville and Keiser. They
reported considerably different profit-maximizing plant populations for the same
hybrid across locations and/or seasons. For example, the profit-maximizing popula-
tion of hybrid PR39W54 was 11.9 (2001), 17.5 (2002), and 18.9 plants/m2 (2003) at
Fayetteville and 10.4 plants/m2 at Keiser (2003).
The aforementioned data are indicative of a positive association between the
environmental yield potential and the required plant population in the field. Norwood
(2001) tested five hybrids at 3, 4.5, and 6 plants/m2 across eight environments (four
seasons, two planting dates) and under rain-fed conditions at the Southwest Research
Extension Center near Garden City, Kansas, USA. He found that the higher popula-
tions usually produce more grain at favorable climatic conditions, but for the stressful
102 I.S. Tokatlidis
Fig. 3 The positive relationship between crop yield potential and the optimum population (i.e.
y = 3.13×−24.2, R2 = 0.67, P < 0.005), regarding the hybrid N4640Bt over three years at four popu-
lations and across six locations [(1) for Ames, (2) for Kanawha, (3) for Nashua, (4) for Sutherland,
(5) for Crawfordsville, and (6) for Lewis] by two row-spacing, i.e. brown dot for 38 cm and blue
dot for 76 cm (Data from Farnham 2001)
season and planting date, average yield decreased from 3,390 to 2,600 kg/ha when
population increased from 3 to 6 plants/m2. When data for the hybrid N4640Bt from
Farnham (2001) are analysed across the location by row spacing combinations, for
10 out of the 12 environments a positive correlation between hybrid crop yield
potential and optimum population is found. According to the linear regression anal-
ysis (r = 0.82, P < 0.001), for a per 1,000 kg/ha increase in yield potential, 3.13 more
plants/m2 are needed for this potential to be fully exploited. It is worth noting the
within-location differences between hybrid optimum population of the inter-row
spacing – 38 cm versus 76 cm – as shown in Fig. 3, emphasizing the strong environ-
ment by population interaction which appears an insurmountable obstacle in deciding
on either hybrid- or site-specific optimum population. Linear regression analysis
between crop yield potential and optimum population over four hybrids and four
N-treatments across 11 growing seasons (Berzsenyi and Tokatlidis 2012), i.e. Fig. 2,
gave a positive correlation (r = 0.67, P < 0.03). Similarly, the 20 crop yield potential–
optimum population pairs from Stanger and Lauer (2006) showed a tendency for
positive correlation (r = 0.57, P < 0.007).
2.2 Implications of Optimum Population Variability on Grain

Yield Productivity and Stability
Interactions among environments, hybrids, and populations create a necessity for

hybrid- and/or site-specific crop management. However, the enormous variation in
yield potential and optimum population across environments mirrors the difficulty
in predicting the most suitable plant population and applying the corresponding
seeding rate. As a consequence, accounting for the divergence from the optimum
population in the degree of hybrid dependence on population, these interactions are
the ultimate cause of substantial yield losses. Table 1 presents the maximum grain
yield loss drawn from reported results, presumably expected at the most stressful
environment if the applied population approaches that suitable for a very favorable
environment, and vice versa.
Data from Blumenthal et al. (2003) obtained under extremely extensive cultiva-
tion are sufficiently explanatory. The authors noted that growers in western Nebraska
are advised to plant for an expected harvest population of 2.72 plants/m2. Indeed, this
population was computed as optimal in the Cheyenne location during 2000. For this
season, however, the quadratic model estimated complete yield loss under a popula-
tion similar to the optimum for the more favorable 1999. On the other hand, farmers
would produce in 1999 under the recommended population of 2.72 plants/m2 35%
less grain compared to the season’s yield potential. Intriguingly, for the Kimball
location, a loss of up to 39% (2000) and 55% (1999) could occur during one of these
two seasons under a population optimal for the other, even though the two seasons
were of the same yield potential (i.e. 1,800 kg/ha). Extremely high potential losses
have been reported for the population-dependent hybrid Norma (Berzsenyi and
Tokatlidis 2012). Total yield destruction could result for seasons like the very dry
1990 in cases where the applied population is high enough to satisfy the require-
ments of favorable seasons similar to 1997.
Such an adverse implication may arise even for less variable environments. In
the study of Shanahan et al. (2004) the low-yielding environment of 1998 exhibited
an optimum population of 5.83 plants/m2, while the high-yielding environment
of the same season gave optimum population of 8.78 plants/m2. Computed maxi-
mum yield losses were 17% and 8.8%, respectively. Regarding the non-Bt group
of hybrids studied by Stanger and Lauer (2006), in the North Central zone the
location of Chippewa Falls exhibited the lowest crop yield potential attainable at
5.78 plants/m2, while yield potential of the most favorable Seymour could be accom-
plished at 11.2 plants/m2. This range of optimum population implies a yield loss up
to 9.3% for Chippewa Falls and up to 17% for Seymour. Seven short-season hybrids
were tested in Romania under dryland conditions, and the over-hybrid optimum
population was 5.38 plants/m2 for the dry 2006 season, fairly approaching that rec-
ommended for the area (Tokatlidis et al. 2011). The next season, however, 9.12
plants/m2 was the required population to take advantage of the adequate rainfalls,
implying a 14% yield loss with the population commonly used in the area. If the
optimum population for the well-rained season was chosen for the dry 2006 season,
the yield loss would reach 27%.
2.3 The Beneficial Role of Plant Yield Potential
Much of the increase in yield per unit area associated with newer hybrids was due
to increased stress tolerance, which allowed growers to adopt higher populations,
104
Table 1 Data from studies concerning two contrasting environments and regarding the crop yield potential (CYP), and optimum population (OP), as well as
the yield loss (YL) whether in each environment a population equal to the optimum of the other was established
CYP OP(q) YL
Hybrid Environment Quadratic equation (kg/ha) (plants/m2) (%) Source
PR3893 Cheyenne, 1999 y = 2,701 + 972.9x −47.2x2 7,720 10.3 35 Blumenthal et al. (2003)
Nebraska (R2 = 0.97)
2000 y = 1,824 + 445.5x−80.6x2 2,440 2.76 100
(R2 = 0.99)
PR3893 Kimball, 1999 y = 731.5 + 487.3x−56.3x2 1,790 4.32 55 Blumenthal et al. (2003)
Nebraska (R2 = 0.94)
2000 y = 1,814−9.98x−35.3x2 1,810 0.14 39
(R2 = 0.98)
PR3860 Anton, Low-yielding y = 1,839 + 1,283x−110x2 5,580 5.83 17 Shanahan et al. (2004)
Colorado, field (R2 = 0.91)
1998 High-yielding y = 2,469 + 1,230x−70x2 7,870 8.78 8.8
field (R2 = 0.92)
non-Bt group North Central Seynour y = 3,020 + 1,560x−70x2 11,710 11.2 17 Stanger and Lauer (2006)
Wisconsin- (R2 = 0.94)
zone Chippewa y = 8,020 + 320x−28x2 8,930 5.78 9.3
Falls (R2 = 0.96)
Short-season Turda, 2006 y = 2,696 + 1,252x−116.3x2 6,070 5.38 27 Tokatlidis et al. (2011)
group Romania (R2 = 0.98)
2007 y = 1,443 + 1,383x −75.82x2 7,750 9.12 14
(R2 = 0.99)
Norma Martonvásár, 1990 y = 491.2 + 615x −67.49x2 1,890 4.56 100 Berzsenyi and Tokatlidis
Hungary (R2 = 0.98) (2012)
1997 y = 3,763 + 1,022x −50.13x2 8,980 10.2 18
(R2 = 0.87)
I.S. Tokatlidis
while the single-plant yield potential has stagnated when any stress is absent (Sangoi
et al. 2002; Duvick 1997, 2005; Tollenaar and Lee 2002; Hammer et al. 2009; Liu
and Tollenaar 2009; Brekke et al. 2011). Consequently, strong hybrid dependence
on population is largely due to the inability of individual plants to exploit larger
input-shares at lower populations. In contrast, the low threshold of the optimum
population could be justified by the ability of individual plants to take advantage of
more resources at lower populations (Fasoula and Fasoula 1997, 2000, 2002;
Tokatlidis et al. 2001, 2011; Fasoula and Tokatlidis 2012).
In order to investigate reliably the impact of plant yield potential on optimum
population and stability, the availability of data regarding hybrids equivalent in crop
yield potential is desired. Although such data are generally scarce, data provided by
Thomison et al. (2011) meet such a presupposition. For four hybrids tested across
5.90–10.4 plants/m2 at S. Charleston, Ohio, crop yield potential values were found
to be almost equal (12,670–13.080 kg/ha). Their optimum population, though, varied
from 6.75 to 10.9 plants/m2 and plant yield potential from 297 to 341 g. These two
measures were inversely correlated (r = −0.94, P < 0.06), reflecting the message that
improved yield at the single-plant level may extend the lower limit of the plant
population range for optimal resource use and crop yield.
Data from Sarlangue et al. (2007) show declining pattern for optimum popula-
tion and increasing pattern for crop yield potential when plant yield potential
increases. They evaluated three hybrids, the short-season Romario, the mid-season
P37P73, and the long-season DK688. Yield response to population was estimated
on the basis of the provided quadratic lines for two seasons, and average values of
plant yield potential, crop yield potential, and optimum population were respec-
tively: 136 g/plant, 9,170 kg/ha, and 13.6 plants/m2 for Romario; 195 g/plant,
10,780 kg/ha, and 12.7 plants/m2 for P37P73; and 267 g/plant, 12,460 kg/ha, and
10.9 plants/m2 for DK688 (Fig. 4). Obviously, improved plant yield potential
influenced crop yield potential favorably and optimum population inversely. These
findings indicate that very high population is not an imperative condition for high
grain yield per unit area to be attained, on the premise that individual plants have the
potential to take advantage of more available resources with less crowding.
Other works similarly report a significant (P < 0.01) negative association between
plant yield potential and optimum population. For example, a significant negative
correlation between plant yield potential and optimum population is computable
from data of Berzsenyi and Lap (2005) including five hybrids across three seasons
(r = −0.59, P < 0.08). Significant negative correlation between plant yield potential
and optimum population was also reported by Berzsenyi and Tokatlidis (2012).
Plant yield potential is obviously crucial for lower optimal populations and thus
a desirable agronomic trait of hybrids sought for variable conditions. When non-Bt
and Bt hybrids of Stanger and Lauer (2006) were scored for plant yield potential
across 10 locations, a positive linear correlation of single-plant yield potential with
crop yield potential was found (r = 0.91, P < 0.001). The finding corroborates a similar
linear relationship found by Tokatlidis (2001). By inference, potential linkage between
these two advantageous attributes determines high yielding population-neutral
hybrids.
106 I.S. Tokatlidis
Fig. 4 Plant yield potential (PYP), crop yield potential (CYP), and optimum population (OP) of
three hybrids, the short-season Romario, the mid-season P37P73 and the long-season DK688 over
2-years, indicate that improved plant yield potential may be combined with improved crop yield
potential and lower optimum population (Data from Sarlangue et al. 2007)
Despite its beneficial agronomic attribute, whether plant yield potential has
improved over time is questionable, and this is partially attributed to stagnation in
harvest index (Duvick 2005; Tollenaar and Lee 2002). On the other hand, high plant
yield potential is associated with high harvest index at lower populations according
to data reported by Hashemi et al. (2005) and Sarlangue et al. (2007). Echarte and
Andrade (2003) found that prolificacy stabilized harvest index and advanced hybrid
reproductive plasticity at low populations. Improved prolificacy was regarded by
Tokatlidis et al. (2005) as the key factor for hybrids of low and broad optimum
population. Boomsma et al. (2009) discovered that strong responsiveness of grain
yield per plant to reduced plant crowding accompanies higher harvest index values
and better nitrogen use efficiency, suggesting a relatively high level of reproductive
partitioning and plasticity. Hence, improvement of yield potential of single plants
is expected to be associated with improved harvest index and prolificacy at lower
populations.
3 Current Crop Management Status
Periodically re-evaluating the response of maize grain yield to population is encour-

aged, since optimum population can differ among hybrids (Widdicombe and Thelen
2002; Stanger and Lauer 2006). However, optimum population has trended upward
over time (Hammer et al. 2009), and thus the economically optimum seeding rate
commonly differs among hybrids (Van Roekel and Coulter 2011). In other words,
the strong reliance of maize crop upon population makes the assessment of hybrid-
and site-specific optimum populations imperative. Nevertheless, under variable climate
conditions the hybrid–population interaction constitutes an almost insurmountable
constraint to deciding on a particular population.
A representative example of the difficulty in addressing the issue of optimal pop-
ulation is the crop production for rain-fed conditions with precipitation commonly
varying across seasons. Norwood (2001) concluded that risk-averse farmers would
probably choose lower populations. He stated that hybrid maturity, plant popula-
tion, and planting date should be diversified for risk management under drought
conditions and to ensure acceptable yields across environments. Blumenthal et al.
(2003) advised growers to plant maize at specific plant population and to increase
above this level only if they are willing to accept the associated risks. Shanahan
et al. (2004) reported results contrasting with the recommendation for hybrid maturity
to be diversified and suggested using variable site-specific management of the plant
population. Even in conventionally irrigated and fertilized maize Popp et al. (2006)
found it hard to reach a particular optimal population.
Planting at lower populations is recommended in drought-prone environments,
where the available resources cannot support a high plant population (Norwood
2001; Blumenthal et al. 2003; Shanahan et al. 2004; Duvick 2005; Tokatlidis et al.
2011; Berzsenyi and Tokatlidis 2012). In addition, special emphasis has been
placed on the time required for a hybrid to reach maturity. It is believed that early-
rather than late-maturing hybrids are more adaptable to lower populations in stressful
seasons, plus they seemingly have potential for avoiding drought (Norwood 2001;
Shanahan et al. 2004; Edwards et al. 2005). However, higher optimum populations
were scored up for shorter- than longer-season hybrids when resources were in
abundance, implying that the short-season hybrids are usually more variable in
optimum population.
The study of Sarlangue et al. (2007) is not supportive of early-maturing hybrids,
since earliness was accompanied by lower yield per area potential and higher opti-
mum population (Fig. 4). A decrease in plant population from 10 to 5 plants/m2
resulted in a 52%, 37%, and 23% decrease in grain yield for the short-season
Romario, the mid-season P37P73, and the long-season DK688, respectively. The
researchers characterized the long-season DK688 hybrid as being more plastic on
account of its ability to explore more resources at low populations. Results from
Shanahan et al. (2004) are not supportive of early-maturing hybrids, alike. They
evaluated two hybrids (the early-maturing PR3860 and the late-maturing PR3752)
at four plant populations (2.5–6.2 plants/m2) across low-, medium-, and high-yielding
environments. Optimum population increased with increasing environmental poten-
tial for the early-maturing hybrid (i.e. 5.83 → 8.78 plants/m2), while it remained
almost unchanged for the late-maturing hybrid (i.e. 6.12 → 6.57 plants/m2).
Berzsenyi and Tokatlidis (2012) reported results with reference to three hybrids
of equal crop yield potential, verifying though stronger dependence on population
for the short- season hybrid and least dependence for the long-season hybrid. A short-
and a full-season hybrid had equivalent asymptotic yield potential as a function of
108 I.S. Tokatlidis
plant population, the threshold of population required to achieve maximum yield,

however, was 19 and 8 plants/m2, respectively (Edwards et al. 2005). Popp et al.
(2006) found that the higher the hybrid maturity, the higher the potential for lower
plant population requirements to achieve required yields.
Avoidance of water deficiency at the crucial stages of grain formation and filling
is of the utmost importance for the crop to withstand the drought stress. The afore-
mentioned studies, though, highlight that the hypothesis of short-season hybrids is
valid only on the premise that population-neutral hybrids are available. However,
accumulated evidence regarding earlier-maturing hybrids supports the population
dependence more so than the population neutrality.
4 Future Crop Management
Under cultivation conditions varying in climate, soil, precipitation, and other con-
stituent parts of agro-ecosystems, the availability of flexible hybrids is imperative.
Hybrids of high yield potential should be able to fully exhibit their potential at a
wide range of low-threshold plant populations. Popp et al. (2006) described as ideal
the high-yielding hybrid at lower populations with grain yield response to popula-
tion following the asymptotic pattern (Fig. 5). Hence, they mirrored a population-
neutral hybrid cultivatable at lower populations and therefore flexible to attain
optimal productivity under either marginal or favorable conditions. Henceforth,
crop management could be characterized by two challenges: the short-term target of
investigation of the less population-dependent among currently cultivated elite
hybrids and the mid-term target of developing population-neutral hybrids.
Fig. 5 The hypothetical example of the ideal maize hybrid exhibits an asymptotic grain yield
response to population. The hybrid with higher yield at lower populations requires lower popula-
tion to attain a given yield (arrow) and has higher profit-maximizing yield (i.e., A vs. B) at lower
populations, mirroring the population-neutral hybrid (Adapted from Popp et al. 2006)
Fig. 6 The grain yield response to plant population of the early to mid-maturity hybrids 34B23
and 34M94, as well as of the full-season hybrids 33G26 and 33J56. Quadratic equations,
optimum population in plants/m2 and crop yield potential in kg/ha are, 34B23:
y = 6,628 + 1,476x−88.89x2 (R2 = 0.84), 8.38, and 12,750; 34M94: y = 3,735 + 2,038x−111.1x2
(R2 = 0.99), 9.17, and 13,080; 33 G26: y = 7,717 + 964.4x−44.44x2 (R2 = 0.96), 10.9, and 12,950;
33J56: y = 9,631 + 900x−66.67x2 (R2 = 0.98), 6.75, and 12,670. Comparison of the full-season
33J56 against the early to mid-season 34M94 show that hybrids equivalent in crop yield poten-
tial are not always of the same value. Data across four populations and over three locations and
three harvest dates from Thomison et al. (2011)
4.1 Seeking Less Population-Dependent Elite Hybrids
Due to population-dependence, the common practice of evaluation of different

hybrids at a single dense stand includes the risk of biased judgment. From this view-
point, the study of Thomison et al. (2011) is exceptionally informative. They evalu-
ated four hybrids, the early to mid-maturity hybrids 34B23 and 34M94 and the
full-season hybrids 33G26 and 33J56, at 5.9, 7.4, 8.9, and 10.4 plants/m2 across
three seasons and three harvest dates at South Charleston, Ohio. They did not find
evident differences in yield among hybrids at the different population levels, imply-
ing hybrids of equivalent value. Consideration of the extended downward quadratic
pattern of the yield to population response for the full-season hybrid 33J56 and the
mid-maturity hybrid 34M94 in particular (Fig. 6) denotes different inferences, nev-
ertheless. For a number of reasons, the first is less reliant on population and thus of
higher value: (1) its optimum population is 26% lower; (2) it satisfies better the
requirements of stressful environments for less plant crowding; (3) when population
varies within the ±30% limits of the optimum, yield loss is up to 2.1%, while the
corresponding yield loss of the second hybrid is 6.2%. However, this desirable attri-
bute of the 33J56 hybrid can be revealed only when hybrid performance is consid-
ered at very low populations. Berzsenyi and Tokatlidis (2012) reported results for
110 I.S. Tokatlidis
two hybrids of similar crop yield potential, the relatively longer-season Maraton and
the shorter-season Norma. For reasons similar to the above, the first exhibited less
dependence on population and was found to be more appropriate for long-term
dryland cultivation, exhibiting the versatility to either adapt to the low-input envi-
ronments or flourish during favorable seasons.
By inference, among currently elite hybrids the less population-dependent have
to be qualified. The justified crucial role of plant yield potential highlights the neces-
sity of evaluating hybrids to target this character. The first option to meet this
demand is to estimate the plant yield potential measure suggested by Yan and
Wallace (1995) in cases where data across a range of populations are already avail-
able. An alternative option is evaluation of the hybrids in the absence of competi-
tion, where plant yield potential is directly measurable. An innovative statistic is
now available, designed for accurate whole-plant field evaluation for high and stable
crop yield and breeding population-neutral cultivars, described in the next section.
Among hybrids already known to perform well at high populations, those which
rank top for plant yield potential are expected to be better suited to variable
situations.
4.2 Development of Population-Neutral Hybrids
Newer hybrids exhibit higher harvest index than older ones when plants are sub-
jected to stresses, thanks to improvement in tolerance to various stresses. On the
other hand, harvest index has not improved when modern hybrids are grown at their
optimal populations or lower (Duvick 2005), on account of grain yield stagnation at
the individual plant level. Hence, improvement of plant yield potential is expected
to be associated with improved harvest index and prolificacy at lower populations,
designating population-neutrality. Indicatively, Echarte and Andrade (2003) and
Luque et al. (2006) reported on parallel improvement in harvest index and plant
yield potential among Argentine hybrids. The necessity of improvement in plant
yield potential was underlined by Duvick (1997, 2005).
Honeycomb breeding was applied at the ultra-low population of 0.74 plants/m2
to target on plant yield potential (Tokatlidis et al. 1998). Forty S5 × S5 hybrids
drawn through single-plant selection within the F2 of the PR3183 hybrid averaged
67% and up to 115% higher plant yield potential than the original hybrid. Six of
these recycled hybrids tested across two locations and two seasons were found to
be considerably less dependent compared to PR3183 (Fig. 7) (Tokatlidis 2001;
Tokatlidis et al. 2001, 2005). According to Duvick (2005) such hybrids are suited
for drought-prone environments, where planting at lower population is prudent
but the ability to utilize occasional higher rainfall by increasing yield per plant
would be desirable.
The innovative honeycomb breeding methodology was established by Prof.
Fasoulas (Tsaftaris 2005). One of the first inviolable principles was selection in
the absence of competition (Fig. 8), meaning that particular emphasis was
placed on plant yield potential from the very beginning (e.g. Fasoulas 1973).
Fig. 7 Direct improvement of plant yield potential though honeycomb breeding led to recycled
hybrids that exhibited less population-dependence compared to the ancestor hybrid PR3183, as it
is shown for six recycled hybrids on average (RC1–6) and a single recycled hybrid (RC5). Data
across two seasons and two locations at three populations from Tokatlidis et al. (2001)
Fig. 8 The primary and inviolable principle of the honeycomb breeding is selection among widely
spaced individual plants that allows them to grow in the absence of competition, thus placing par-
ticular emphasis on the determinant factor of population-neutrality, i.e. the plant yield potential
Since then, further advancement in the methodology, and particularly the thor-
ough explanation of the suspending role of competition in breeding (Fasoulas
1973, 1988, 1993; Fasoulas and Fasoula 1995; Fasoula and Fasoula 1997), has
opened the possibility of even more progress through honeycomb breeding.
112 I.S. Tokatlidis
To couple in parallel performance in the absence of competition and farming

conditions, whole-genome phenotype is accomplished by partitioning the crop
yield potential into three components, which are plant yield potential, tolerance
to stresses, and responsiveness to inputs (Fasoula and Fasoula 2000, 2002).
Fasoula (2008) incorporated the aforementioned crop yield potential compo-
nents into two equations to evaluate individual plants and entries (i.e. progeny
lines or cultivars), while Fasoula and Tokatlidis (2012) analyzed the major
principles of the method to develop cultivars which use resources effectively,
tolerate biotic and abiotic adversities, and are capable of broad adaptation.
Briefly, the equation A assesses the crop yield potential of individual plants
(PCYP), and the equation B the crop yield potential of the entries (ECYP),
enabling the breeder to apply single-plant selection based on equation A within
lines singled out through equation B. Evaluation according to these equations is
reliable only when individual plants and entries are assessed in the absence of
the confounding effects of competition and soil heterogeneity, which is achieved
with the utilization of the honeycomb selection designs that enable the applica-
tion of ultra-high selection pressures. Each equation consists of two parameters;
the first measures the plant yield potential in equation A and entry yield poten-
tial in equation B, while the second, which is common in both equations, mea-
sures the entry’s stability of performance. More specifically, single-plant yield
( x) is expressed as its ratio to the average yield of the surrounding plants within
a ring of a chosen size ( xr ) . The square of this ratio, ( x / xr ) 2 , called the
coefficient of plant yield, devoid of the masking effect of soil heterogeneity,
allows plants to be ranked objectively according to their true yield potential.
Stability of the entry to which each plant belongs is measured by the coefficient
of homeostasis ( x / s ) 2 , where x and s are the mean and the standard deviation,
respectively. The product of the two coefficients, PCYP = ( x / xr ) 2 ·( x / s ) 2 ,
represents the crop yield potential at the plant level. Entry crop yield potential,
ECYP = ( x / xt ) 2 ·( x / s ) 2 , comprises the coefficient of entry yield, ( x / xt ) 2 ,
where xt is the overall mean in the trial, and the coefficient of homeostasis,
( x / s)2 .
Reliability of the ECYP to assess the value of hybrids was investigated by
Tokatlidis et al. (2011). Their results justified the suitability of the population-
neutral hybrids for drought-prone environments as opposed to the dependent
ones, as well as the ability to anticipate hybrid performance at crop populations
from evaluation in the absence of competition. Hence, to combat the challenge
of climatic changes the honeycomb breeding methodology constitutes a unique
breeding tool to meet the demand for population-neutral hybrids. Apart from a
number of studies which are supportive of the method in maize and other crops
(Fasoula and Tokatlidis 2012), regarding the suggested equations Vlachostergios
et al. (2011) found that PCYP was an effective selection tool for organic breed-
ing in lentils, while Papadopoulos and Tokatlidis (2011) qualified the ECYP as
a stability criterion in dry beans and classified it according to the agronomic
concept of stability, i.e. to designate cultivars performance in accordance with
the available inputs.
5 Synopsis
Future maize producers may have to make crop management decisions for highly
diverse agro-ecosystems, due to changing climate scenarios. To meet the require-
ment for effective resource use and optimize grain productivity, availability of plastic
hybrids characterized by general adaptability in the matter of population is impera-
tive. Feasibility of this target, however, is complicated by the hybrid–population
interaction.
Hybrid response to spatial and temporal heterogeneity differs among hybrids
(Williams et al. 2008). Hence, in diverse environments the hybrid–population inter-
action is commonly strong, designating population-dependence. Indicatively, the
optimum population has increased throughout the hybrid era, while hybrids differ in
optimum population (Duvick 1997, 2005; Tollenaar and Lee 2002; Widdicombe
and Thelen 2002; Boomsma et al. 2009; Brekke et al. 2011). Intriguingly, the profit-
maximizing population of a hybrid may differ from the yield-maximizing one (Popp
et al. 2006).
Population dependence, though, is a major barrier to reaching a site- and hybrid-
specific decision on optimal population. Researchers working under greatly variable
conditions found it hard to recommend a population (Norwood 2001; Blumenthal
et al. 2003; Shanahan et al. 2004; Berzsenyi and Tokatlidis 2012). Such a problem is
detectable even in studies under less variable environments (Farnham 2001; Popp
et al. 2006; Stanger and Lauer 2006; Boomsma et al. 2009; Tokatlidis et al. 2011).
Maize hybrid dependence on population adversely affects crop stability in grain
yield due to a number of causes, e.g. occasional missing plants, deteriorated stand
uniformity, inefficient resource use, increased lodging, asynchronous flowering,
and increased barrenness (Tokatlidis and Koutroubas 2004). Furthermore, when
farmers grow maize under variable climatic condition, enormous variability in opti-
mum population may cause substantial grain yield and income penalty. Several
relevant studies provide data supporting such an implication (Norwood 2001;
Blumenthal et al. 2003; Shanahan et al. 2004; Stanger and Lauer 2006; Boomsma
et al. 2009; Tokatlidis et al. 2011; Berzsenyi and Tokatlidis 2012).
Important direct effects of the ongoing global weather changes on crop yields
will be through changes in temperature, precipitation, length of growing season, and
timing of extreme or critical threshold events relative to crop development
(Southworth et al. 2000; Cutforth et al. 2007; Tingem et al. 2009; Hatfield et al.
2011). Because of these events crops might encounter more sharply fluctuating
environmental situations in the future than currently occur. Vulnerability of agricul-
tural systems to climate variance might reasonably be more severe in crops that
interact strongly with population. In turn, maize instability would probably worsen
and thus sustainability of the crop under such circumstances would be in danger.
Stagnation in plant yield potential alone accounts for severe maize hybrid
dependence on high populations. Differences in grain yield per unit area through-
out the hybrid era have been shown to be a function of population rather than yield
potential per se (Duvick 1997, 2005; Tollenaar and Lee 2002; Hammer et al. 2009;
114 I.S. Tokatlidis
Brekke et al. 2011; Van Roekel and Coulter 2011). Results from Thomison et al.
(2011), however, indicate that high yield per area is not solely accomplishable at
very high populations on the condition that the hybrid has improved plant yield
potential (Fig. 6). Other studies (Norwood 2001; Sarlangue et al. 2007; Berzsenyi
and Lap 2005; Tokatlidis et al. 2011; Berzsenyi and Tokatlidis 2012) verify that
improved plant yield potential is a determinant of adaptation to low plant popula-
tions. Improved plant yield potential coupled with high crop yield potential can
exist (Tokatlidis 2001; Luque et al. 2006; Popp et al. 2006). Hence, hybrids which
combine both these attributes and tolerate high populations are designated popula-
tion-neutral hybrids that accomplish their crop yield potential at a wide population
range.
Planting at lower populations when the available resources cannot support a high
plant population is an inevitable option (Norwood 2001; Blumenthal et al. 2003;
Shanahan et al. 2004; Duvick 2005; Tokatlidis et al. 2011; Berzsenyi and Tokatlidis
2012). On the other hand, earliness in maturity is considered a drought-avoidance
mechanism at the critical grain filling stage (Norwood 2001; Shanahan et al. 2004;
Edwards et al. 2005). In the matter of maturity, however, relevant studies provide
data showing that, in general, short-season hybrids may require very high popula-
tions to optimize resource use when high rainfalls occur (Norwood 2001; Shanahan
et al. 2004; Edwards et al. 2005; Popp et al. 2006; Sarlangue et al. 2007; Berzsenyi
and Tokatlidis 2012), implying for producers considerable yield and income loss at
favorable seasons. Consequently, hybrid maturity alone cannot address the major
challenge of adapting maize crop to conditions of great variance. Population-neutral
hybrids of high plant yield potential instead, could yield sufficiently when avail-
ability of resources would not support large plant numbers per unit area and, in
parallel, they could take advantage when resources are in abundance.
Data from Shanahan et al. (2004) as well as from Thomison et al. (2011) (Fig. 6)
are indicative that seeking among elite hybrids those which are less dependent on
population is sound advice. Berzsenyi and Tokatlidis (2012) set the issue as a pri-
mary hypothesis and managed to discover a particular hybrid that was substantially
the most suitable for extremely diverse seasons. Apparently, improved plant yield
potential is a determinant of population neutrality. In turn, the estimation of this
constituent element of crop yield potential in hybrids allocated for cultivation is a
major challenge for agronomists.
Obviously, breeding of population-neutral hybrids is a fundamental issue for
future agriculture. Tokatlidis et al. (2001) opened the possibility of obtaining such
hybrids (Fig. 7) through honeycomb breeding targeted directly at plant yield poten-
tial (Tokatlidis et al. 1998). Advanced selection criteria were suggested by Fasoula
and Tokatlidis (2012) to create new cultivars with the potential to exploit effectively
even marginal environments and meet the needs of sustainable agricultural systems.
They speculated that since the method places particular emphasis on selection at
ultra-low population to counteract the disturbing effects of plant-to-plant inference
in equal share of inputs, it substantially improves plant yield potential, thereby
expanding the lower limit of the optimum population. On the other hand, selection
criteria consider the genotype ability to withstand stresses, and thus new cultivars
are able to perform well at high populations. The unique qualities of population-neutral
hybrids guarantee their advantageous position to adapt well to climatic and other
environmental changes and promote sustainable agricultural systems.
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Intercropping
Ehsan Neamatollahi, Mohammad Reza Jahansuz, Dariush Mazaheri,

and Mohammad Bannayan
Abstract Intensive agricultural systems have negative impacts on soil and water
quality, and on biodiversity conservation. The evolution of intercropping is con-
trolled by a high degree of biodiversity. In sharp contrast intensive agricultural sys-
tems uses monocultures associated with high input of chemical fertilisers and
pesticides. Intercropping involves two or more crops in the same field at the same
time. Intercropping increases biodiversity, improving the ability of an agricultural
system to handle changes in growing conditions. Intercropping plays a pivotal
role for increasing land use efficiency, weed suppression, enhanced ecological
services and greater economic profitability. Benefits of intercropping include
improved yields and yield stability, enhanced use of water and nutrients, increased
weed suppression, increased pest and disease resistance, reduced soil erosion and
improved forage quality.
This review reports the relationships between plants in intercropping to under-
stand and design intercropping systems, with special focus on intercropping
efficiency. The most common index showing yield advantage of intercropping is the
land equivalent ratio (LER). The LER provides a standardized basis so that crops
can be added to form ‘combined’ yields. The LER indicates competitive effects by
comparison of individual LER. LER can measure relative yield advantage. We also
present new ways to determine and use the LER.
Keywords Intercropping • Mazaheri’s LER table • Intercropping efficiency

• Unilateral cooperation • Unilateral inhibition
E. Neamatollahi (*) • M.R. Jahansuz • D. Mazaheri

Department of Agronomy, University of Tehran, P.O. Box. 4111, Karaj, Iran
e-mail: neamatollahi_e@ut.ac.ir
M. Bannayan
Department of Agronomy, Ferdowsi University of Mashhad,
P.O. Box. 91775-1163, Mashhad, Iran
120 E. Neamatollahi et al.
1 Introduction
In developed regions of the world, crops are primarily grown as sole crops and
monocultures. However, increasing awareness of the link between agricultural
practice, environmental issues and long-term stability of existing food production
systems has put focus on the role that greater crop diversity in time, crop rotation
and space, intercropping may reduce the extent of these problems. Intercropping
represents one way of increasing crop diversity.
As a result of differences in the way component crops respond to and affect the
environment in which they are grown, intercrops may use available growth resources,
light, water and nutrients, more efficiently, reduce the prevalence of disease and
pests and reduce weed infestation compared to sole crops. The greatest intercrop
advantages are attained when the species that are mixed, differ markedly either
morphologically, phenologically or physiologically.
In recent years, there has been increased interest in agricultural production sys-
tems in order to achieve high productivity and promote sustainability over time.
From ancient times, farmers developed different cropping systems to increase pro-
ductivity and sustainability; they included crop rotation, relay cropping, and inter-
cropping of annual cereals with legumes. Intercropping has gained interest because
of potential advantages it offers over yielding, improved utilization of growth
resources by the crops and improved reliability from season to season. Harsh envi-
ronmental conditions under Africa and many areas of Asia (Banik et al. 2006),
which is detrimental to crop plants, may be congenial for weeds, the menacing pest
in crop production. Use of herbicides in any crop mixture is a risky endeavor and
not an eco-friendly approach. Biological and cultural weed controls are important
components of Integrated Weed Management. Researchers are confronted with the
complex problem of weed management by ecological means, giving due consider-
ation to minimal use of chemicals with least disturbance to the environment. Weed
management in intercropping, however, has hardly been studied to date (Altieri and
Liebman 1986; Moody and Shetty 1979; Midya et al. 2005).
Intercropping is an old and widespread practice used in low input cropping sys-
tems in many areas of the world (Anil et al. 1998). During the twentieth century
there was a shift from mainly labour-intensive systems to more optimised cropping
through the use of external inputs, especially synthetic fertilisers and pesticides
(Crews and Peoples 2004). A growing interest in intercropping systems has been
initiated in developed countries due to the increasing awareness of environmental
degradation arising from the heavy use of non-renewable resources (Fujita et al.
1992). Intercrops can use the available environmental resources more efficiently
and thus result in higher yields than monocrops (Vandermeer 1990) (Fig. 1).
The reasons for the higher yield in such systems is that the intercropped species
do not compete for exactly the same growth resource niche and thereby tend to use
the available resources in a complementary way (Hauggaard-Nielsen et al. 2001a,
2003). Competition is one of the factors that can have a significant impact on yield
of crop mixture compared with pure cereal stands (Caballero et al. 1995). Higher
yields have been reported when competition between the two species of the mixture
Intercropping 121
Fig. 1 Agroforestry is a form of intercropping. Agroforestry is an integrated approach using the

interactive benefits from combining trees and shrubs with crops or livestock. Agroforestry com-
bines agricultural and forestry technologies to create more diverse, productive, profitable, healthy,
and sustainable land-use systems. Here tree and wheat intercropping
was lower than competition within the same species (Vandermeer 1990). Interplant
competition usually includes competition for soil water, available nutrients, and
solar radiation (Buxton and Fales 1993).
Competition can also have a significant impact on the growth rate of the different
species used in mixtures. A need for yield stability and limited economic risks is often
the reason for using more diversified cropping strategies in developing countries
(Vandermeer 1989). Most farming strategies in Europe though, have over the past 3–4
decades, implemented a great use of chemical fertilizers, pesticides and mechanization
that has created production of sole crops and monocultures (Crews and Peoples 2005).
However, the political agenda favors the diffusion of alternative techniques to reduce
the environmental impact by reducing synthetic inputs in the farming system. In crop-
ping systems, some services can be immediately identified. One is the restoration of
soil fertility provided to shifting cultivators during regrowth of natural vegetation after
a period of cropping; at the same time yield reducing weeds are suppressed.
Another service provided to farmers is the supplementation of soil nitrogen by
leguminous species and their symbionts; this benefits non-leguminous crops that
follow them. Yet another service, seen in rotations and crop mixtures, is the protec-
tion of plants of individual crop species from their host-specific predators and dis-
ease organisms. Populations of such crop enemies tend to explode when they find
pure stands offering uniformly accessible food. Each of these services has its own
dynamics, requiring special conditions and more or less time. The effects of diver-
sity in cropping systems on pest and disease attack have been considered elsewhere
(Vandermeer 1989; Trenbath 1993).
Intensive agricultural systems are often based on optimising the productivity of
monocultures. In those systems, crop diversity is reduced to one or very few species
that are generally genetically homogeneous, the planting layout is uniform and
symmetrical, and external inputs are often supplied in large quantities. Such
Table 1 Definitions of intercropping

Researcher Definition
Willey (1979a) The growing of two or more crops simultaneously
on the same area of ground where they are simultaneous
for significant part of their growing periods
Beets (1982) Growing more than one crop on the same piece of land
during one calender year
Andrews and Kassam (1976) The intensification of cropping in time and space
Francis (1986) dimensions. Growing two or more crops on
the same field in the year
Vandermeer (1989) Growing two or more crops on the same field any year
Stinner and Blair (1990) Using the same field to produce two or more crops a year
systems are widely criticised today for their negative environmental impacts, such
as soil erosion and degradation, chemical contamination, loss of biodiversity and
fossil fuel use (Giller et al. 1997; Griffon 1999; Tilman et al. 2002).
Conversely, multispecies cropping systems may often be considered as a practical
application of ecological principles based on biodiversity, plant interactions and other
natural regulation mechanisms. They are assumed to have potential advantages in pro-
ductivity, stability of outputs, resilience to disruption and ecological sustainability,
although they are sometimes considered harder to manage (Vandermeer 1989). Yield
increments resulting from mixed intercropping (Jensen 1996b; Anil et al. 1998; Dapaah
et al. 2003; Chen et al. 2004) were attributed mainly to the presence of complimentary
effects, better resource use efficiency of the mixed cultures and the buffering effects of
the mixtures against diseases and weeds (Willey 1979b; Anil et al. 1998) (Table 1).
2 Main Intercropping Types
Intercropping is the growing of two or more crop species in the same field during a
growing season (Ofori and Stern 1987). A large body of literature exists on inter-
cropping, a considerable part of which deals with studies carried out in Africa, Asia
and Latin America (Connolly et al. 2001; Vandermeer 1989). Basic to all intercrop
studies is the assumption that some advantage(s) is achieved from mixing crops
opposed to growing them alone and that these advantages are the result of differ-
ences in the way species exploit or act in relation to the environment in which they
are grown. Andrews and Kassam (1976) identified four main intercrop types.
2.1 Mixed Intercropping
Mixed intercropping is growing two or more crops simultaneously with no distinct

row arrangement (Andrews and Kassam 1976) Broadcast or random establishment
as in indigenous slash and burn or fallow agriculture and current industrial grain
Intercropping 123
Fig. 2 Mixed intercropping is growing two or more crops simultaneously with no distinct row
arrangement. Here cereal and grain legumes intercropping
mixtures (Gomez and Gomez 1983). It is also termed mixed cropping with a similar
description (Andrews and Kassam 1976; Beets 1982) but with the earlier mentioned
potential confusion. Willey (1979a) adds the point that there can also be rows with
mixing within the row that is still termed mixed intercropping. Mixed intercrops are
typically planted together but may have different crop maturation times. Obviously
this type of intercropping will have the most intense interactions between crop spe-
cies or varieties (Fig. 2).
2.2 Row Intercropping
The growing of two or more crops simultaneously where one or more crops are
planted in rows (Andrews and Kassam 1976). Vandermeer (1990) observes that this
is the pattern of intercropping usually encountered in intensive agriculture, such as
peas with canola, alternating maize and soybeans and various tree-based systems.
Beets (1982) add that it occurs ‘in a fixed pattern of spacing and rows’. Many tradi-
tional polyculturalists make use of this method (Fig. 3).
2.3 Strip Intercropping
The growing of two or more crops simultaneously in strips wide enough to permit
independent cultivation but narrow enough for the crops to interact agronomically
(Andrews and Kassam 1976 ) . Vandermeer ( 1990 ) observed that this form of
Fig. 3 Row intercropping is the growth of two or more crops simultaneously where one or more
crops are planted in rows. Here Kura claver and corn intercropping
intercropping is more common in highly mechanised systems. Clearly, as cropping

is increasingly mechanised, the distance between intercrops is increased and
their potential level of interaction is decreased. This is an important consider-
ation later in the discussion of the nature, principles and benefits of crop compe-
tition and complementarily in intercrops (Fig. 4).
2.4 Relay Intercropping
The growing of two or more crops simultaneously during part of the life cycle of
each is defined as relay intercropping. A second crop is planted after the first crop
has reached its reproductive stage of growth, but before it is ready for harvest
(Andrews and Kassam 1976). Stinner and Blair (1990) added that there must be
some overlap in the life cycles of the two crops (otherwise it would be termed
sequential cropping). Andrews and Kassam (1976) referred to this as relay planting
and added that the following crop can be seeds or seedlings. Some researchers
believe that a distinguishing feature of relay intercropping is that the flowering
periods of the two crops overlap- the most stringent definition, but one relevant to
crop interactions (Figs. 5 and 6).
Intercropping 125
Fig. 4 Strip intercropping is the growth of two or more crops simultaneously in strips wide enough
to permit independent cultivation but narrow enough for the crops to interact. Here corn and
soybean intercropping
Fig. 5 Relay intercropping is the growth of two or more crops simultaneously during part of the
life cycle of each is defined as relay intercropping. Here cereal rye and hairy vetch intercropping
Strip row
Mixed relay
cassava other crops
Fig. 6 Intercropping types, e.g. cassava and other crops intercropping
This review paper shows relationship between plants in intercropping for better
understanding and designing of intercropping systems with attention to the methods
of evaluate intercropping efficiency.
3 Legume and Cereal Intercropping
When a legume is grown in association with another crop, intercropping, com-

monly a cereal, and the nitrogen nutrition of the associated crop may be improved
by direct nitrogen transfer from the legume to cereal (Giller and Wilson 1991).
Legumes, with their adaptability to different cropping patterns and their ability
to fix nitrogen, may offer opportunities to sustain increased productivity (Jeyabal
and Kuppuswamy 2001). Therefore, productivity normally is potentially
enhanced by the inclusion of a legume in a cropping system (Maingi et al. 2001).
Legume intercrops are also potential sources of plant nutrients that comple-
ment/supplement inorganic fertilizers (Banik and Bagchi 1994; Ofori and Stern
1987).
In addition, legume intercrops are included in cropping systems because they
reduce soil erosion (Giller and Cadisch 1995) and suppress weeds (Exner and
Cruse 1993). Intercropping systems, especially those employing cereals with
Intercropping 127
legumes, have several major advantages such as higher total yield and better land
use efficiency (Dhima et al. 2007) yield stability of the cropping system (Lithourgidis
et al. 2006) better utilization of light, water, and nutrients (Javanmard et al. 2009)
improved soil conservation (Anil et al. 1998) and better control of pests and weeds
(Banik et al. 2006; Vasilakoglou et al. 2008).
Grass–legume intercrops are common in natural ecosystems, but now are rarely
used in developed countries, except for certain intercropping systems used for ani-
mal feed. A renewed interest in intercropping (Malézieux et al. 2009) and particu-
larly in intercrops of cereals with legumes has risen again lately. In particular,
cereals seem to be more competitive for soil inorganic nitrogen (N) than legumes
due to faster and deeper root growth and higher demand in N (Fujita et al. 1992;
Hauggaard-Nielsen et al. 2001b, 2003).
Consequently, the legumes usually increase their reliance on symbiotic N2
fixation (Li et al. 2006). The complementary use of N sources between species
could be of particular interest in low-N-input cropping systems and organic farm-
ing, especially for cereals with high N requirements such as durum wheat. The dif-
ferent legumes used in the intercropping systems such as pea and common vetch are
often less competitive than cereals and thus may require higher planting densities
than cereals to produce intercropping benefits.
Izaurralde et al. (1990) used intercrops of pea with barley in different planting
densities and found that grain, straw and dry matter yields of the mixtures increased
with increasing pea planting density. Other experiments using different seeding
ratio of pea intercropped with barley and oat showed no significant dry matter or
N yield benefits as the intercrops were heavily dominated by the cereal component
because of the high cereal densities (Carr et al. 1998). Intercropping of cereals with
legumes has been a common cropping system in rain-fed areas and especially in the
Mediterranean countries (Papastylianou 1990; Anil et al. 1998; Lithourgidis et al.
2004, 2006). Intercropping of cereals with legumes improves soil conservation
(Anil et al. 1998), favors weed control (Vasilakoglou et al. 2005; Banik et al. 2006),
provides better lodging resistance (Anil et al. 1998), yield stability (Lithourgidis
et al. 2006), hay curing, and forage preservation over pure legumes and may increase
crude protein percentage, protein yield, and length of optimum harvest period over
grasses (Qamar et al. 1999).
Several factors can affect growth of the species used in intercropping, including
cultivar selection, seeding ratios, and competition between mixture components
(Droushiotis 1989; Roberts et al. 1989; Papastylianou 1990; Caballero et al. 1995;
Carr et al. 2004).
In the Mediterranean countries, one of the legumes extensively used in intercrop-
ping with cereals is common vetch, an annual legume with a climbing growth habit
and high levels of protein (Thomson et al. 1990; Anil et al. 1998). A number of dif-
ferent cereals have been proposed to be appropriate for intercropping with common
vetch such as barley, oat, triticale, wheat (Caballero and Goicoechea 1986; Thompson
et al. 1992; Lithourgidis et al. 2006). Intercropping of grain legumes with cereals
might be a method to improve the crops utilization of soil mineral N to improve
yield (Hauggaard-Nielsen et al. 2001a; Jensen 1996b), and grain quality (Gooding
et al. 2007), and at the same time minimizing N emissions during autumn and winter
(Hauggaard-Nielsen et al. 2003; Jensen 1996a).
Intercropping combined with catch crop and spring cereal cropping are strat-
egies for potentially enhancing yield stability due to better use of natural
resources. Intercropping of cereals and legumes is important for the develop-
ment of sustainable food production systems, particularly in cropping systems
with limited external inputs (Bulson et al. 1997; Dapaah et al. 2003). In the
tropics, cereal/legume intercropping is commonly practiced because of yield
advantages, greater yield stability and lower risks of crop failure that are often
associated with monoculture (Jensen 1996b; Hauggaard-Nielsen et al. 2001a;
Tsubo et al. 2005).
One of the advantages of a cereal/legume intercropping in a humid tropical envi-
ronment is the component crops ability to utilize different sources of N (Willey
1979b; Benites et al. 1993). The cereal may be more competitive than the legume
for soil mineral N, but the legume can fix N symbiotically if effective strains of
Rhizobium are present in the soil.
4 Evaluate Intercropping Efficiency
Several indices such as land equivalent ratio, relative crowding coefficient, competi-
tive ratio, aggressivity, actual yield loss, monetary advantage, and intercropping
advantage have been developed to describe the competition and the economic
advantage in intercropping (Banik et al. 2000; Ghosh 2004; Midya et al. 2005).
Indices can express various attributes of competition in plant communities, includ-
ing competition intensity, competitive effects, and the outcome of competition.
They help in the interpretation of complex data and allow comparison of results
from different studies with the use of the same index.
The most popular single index for expressing the yield advantage of intercrop-
ping systems is probably the LER, defined as the relative land area that is required
for monocrops to produce the same yields as intercrops. The advantages of the LER
are that (a) it provides a standardized basis so that crops can be added to form ‘com-
bined’ yields, (b) it can indicate competitive effects through comparison between
individual LERs, and (c) it can be taken as a measure of the relative yield advantage
(Mead and Willey 1980).
Similarly, the relative crowding coefficient is used to quantify the effects of com-
petition. Additional indices of the intensity of competition, aggressivity, competi-
tive ratio, involve measures of plant biomass in monocultures and mixtures in more
complex formulae. Economic feasibility of intercropping systems is ultimately
determined by their monetary advantage. Often, a biologically efficient system is
not economically viable and cannot be recommended for wide scale adoption.
Competition indices usually do not provide any information of the economic advan-
tage of intercropping systems and thus a number of indices, the monetary advantage
index and the intercropping advantage, are used together to assess the economic
viability of intercropping systems.
Intercropping 129
The advantage of intercropping and the effect of competition between the two
species used in a mixture were calculated using different competition indices as follows:
4.1 Relative Yield Total (RYT)
To analyze the crop performance of the intercropping system, the relative yield total
(RYT) was calculated for all replacement series of the simulated crop stands (DeWit
and Vanden Bergh 1960):
Ylc Ycl
RYT = +
Yll Ycc
Where Y is the crop yield (kg ha−1) and suffixes L and C denote leek and celery,
respectively. Thus, Ylc is the yield of leek when grown in mixture and Yll is the yield
of leek when grown in monoculture. Ycl and Ycc are the corresponding yields for
celery in mixture and monoculture, respectively.
4.2 Land Equivalent Ratio (LER)
The land equivalent ratio (LER) was used as the criterion for mixed stand advantage
as both pea and cereal were desired species in the mixtures. In particular, LER
indicates the efficiency of intercropping for using the environmental resources com-
pared with monocropping. The value of unity is considered the critical value for this
index. When LER is greater than one the intercropping favors the growth and yield
of the intercropped species, whereas when LER is lower than one the intercropping
negatively affects the growth and yield of the species (Mead and Willey 1980). The
LER was calculated as:
LER = (LERp + LERc),
Ypi
LERp =
Yp
Yci
LERc =
Yc
Where Yp and Yc are the yields of pea and cereal, respectively, as monocrops and Ypi
and Yci are the yields of pea and cereal, respectively, as intercrops.
4.3 Land Equivalent Coefficient (LEC)
′′
LEC = La * Lb ′′
Where, La = LER of main crop and Lb = LER of intercrop (Anil et al. 1998).
4.4 Relative Crowding Coefficient (K)
The relative crowding coefficient (K) is a measure of the relative dominance of one
species over the other in a mixture (Ghosh 2004). The relative crowding coefficient
was calculated as:
K = Kp.Kc
Ypi Zci
Kp =
(Yp-Ypi) Zpi
Yci Zpi
Kc =
(Yc-Yci) Zci
Where Zpi is the sown proportion of pea in mixture and Zci the sown proportion of
cereal in mixture. When relative crowding coefficient is greater than one the species
is more competitive, when relative crowding coefficient is equal to one there is no
competition, and when it is lower than one the species is less efficient in resource
use resulting in relative yield loss.
4.5 Aggressivity (A)
Aggressivity is often used to indicate how much the relative yield increase in ‘a’
crop is greater than that of ‘b’ crop in intercropping (Agegnehu et al. 2006). The
aggressivity is derived from the equation:
⎛ Yci ⎞ ⎛ Ypi ⎞
Ac = ⎜ ⎟ -⎜ ⎟
⎝ Yc Zci ⎠ ⎝ Yp Zpi ⎠
⎛ Ypi ⎞ ⎛ Yci ⎞
Ap = ⎜ ⎟ -⎜ ⎟
⎝ Yp Zpi ⎠ ⎝ Yc Zci ⎠
if Ac = 0, both crops are equally competitive, if Ac is positive then the cereal spe-
cies is dominant, if Ac is negative then the cereal species is the dominated
species.
4.6 Competitive Ratio (CR)
Competitive ratio (CR) is another way to assess the competition between different
species. The competitive ratio gives a better measurement of competitive ability of
the crops and also is more advantageous as an index over relative crowding
coefficient and aggressivity. The competitive ratio represents simply the ratio of
individual LERs of the two component crops and takes into account the proportion
Intercropping 131
of the crops in which they are initially sown. The competitive ratio is calculated
according to the following formula (Dhima et al. 2007):
⎛ LERp ⎞ ⎛ Zci ⎞
CRp = ⎜ ⎟⎜ ⎟
⎝ LERc ⎠ ⎝ Zpi ⎠
⎛ LERc ⎞ ⎛ Zpi ⎞
CRc = ⎜ ⎟⎜ ⎟
⎝ LERp ⎠ ⎝ Zci ⎠
4.7 System Productivity Index (SPI)
Another index for assessing intercrops is the system productivity index (SPI), which
standardizes the yield of the secondary crop (cereal) in terms of the primary crop
(pea) (Agegnehu et al. 2006) and is calculated as:
⎛ Sp ⎞
SPI = ⎜ ⎟ Yc + Yp
⎝ Sc ⎠
Where Sc and Sp are the mean yield of cereal and pea in monocrop and Yc and Yp are
the mean yield of cereal and pea in mixed culture.
4.8 Actual Yield Lose or Gain (AYL)
Moreover, Banik et al. (2000), reported that the actual yield loss (AYL) index, based
on yield per plant, gave more precise information than other indices about the com-
petition between and within component crops and the behavior of each species in
intercropping. The actual yield loss is the proportionate yield loss or gain of inter-
crops compared with the respective monocrop, i.e. it takes into account the actual
sown proportion of the component crops with their pure stand. In addition, partial
actual yield loss (AYLp or AYLc) represent the proportionate yield loss or gain of
each species when grown as intercrops relative to their yield in pure stand. The actual
yield loss is calculated according to the following formula (Banik et al. 2000):
AYL = AYLp + AYLc
⎡ (Ypi / Zpi) ⎤
AYLp = ⎢ ⎥ -1
⎣ (Yp / Zp) ⎦
⎡ (Yci / Zci) ⎤
AYLc = ⎢ ⎥ -1
⎣ (Yc / Zc) ⎦
The actual yield loss can have positive or negative values indicating an advantage or
disadvantage accrued in intercrops when the main objective is to compare yield on
per plant basis.
4.9 Monetary Advantage Index (MAI) and Intercropping

Advantage Index (IA)
None of the abovementioned competition indices provides any information about

the economic advantage of an intercropping system. For this reason, the monetary
advantage index (MAI) and the intercropping advantage (IA) index were calculated
according to Banik et al. (2000), and Ghosh (2004) as follows:
MAI = Valueof combined intercrops * LER − 1

LER
IA = IAc + IAp
IAc = AYLc Pc
IAp = AYLp Pp
Pp is the commercial value of pea silage (the current price is $42 per Mg), and Pc is
the commercial value of cereal silage (the current price is $31 per Mg). Value of
combined intercrops was calculated as: YpiPp + YciPc.
4.10 Area Time Equivalency Ratio (ATER)
(Rya * ta) + (Ryb * tb)

ATER =
T
Where, Ry = Relative yield of species ‘a’ or ‘b’ i.e., yield of intercrop/yield of main
crop, t = duration (days) for species ‘a’ or ‘b’ and T = duration (days) of the inter-
cropping system (Hiebsch and McCollum 1987).
4.11 Area Harvest Equivalent Ratio (AHER)
Balasubramanian and Sekayange (1990) proposed a concept called area harvest

equivalency ratio, AHER, as nearer to the true value of intercrop yield advantage.
Ayi Byi
AHER = +
n.Ays n.Bys
Where, Ayi and Byi are A, B yield in intercropping, respectively and n is number of
harvest. For example in alfalfa can be harvested four or five period and ays and bys
are a,b yield in sole crop.
The most popular single index for expressing the yield advantage of intercrop-
ping systems is probably the land equivalent ratio and the advantages of the land
Intercropping 133
Table 2 Mazaheri’s LER table; Evaluate land equivalent ratio, suitable and being dominant or not
1.8
1.6
1.4
1.2
LERa 1 LER > 1
0.8
0.6
0.4 LER < 1
0.2
0
0 0.2 0.4 0.6 0.8 1 1.2 1.4 1.6 1.8
LERb
equivalent ratio are that, and then we innovation a way for determine the land equiv-
alent ratio with table.
5 Evaluate Land Equivalent Ratio with Table: Mazaheri’s

Land Equivalent Ratio Table
In this table we can evaluate land equivalent ratio, suitable and being dominant or
not. That LERa is the first plant land equivalent ratio and LERb is the second plant
land equivalent ratio at intercropping (Table 2).
6 Density
In terms of density, intercropping systems range from replacement, plants of a com-

ponent crop in sole cropping are replaced with those of another to give the same
final density, to additive, where plant density and the arrangement of intercrops are
the same as in sole cropping, with the final plant density being the sum of the densi-
ties of the sole crops. In terms of components, intercrops can be combinations of
annuals, maize-legume, perennials including trees or a mixture of the two usually
referred to as agroforestry.
6.1 No Competition (Neutralism)
In this case, intercropping yield is equal with mono cropping yield and then farmer
has not relish for use of inter cropping (Inter specific competition = Intra specific
competition) (Fig. 7).
A and B intercropping yield
A plant B plant
A plant density
%0 25 50 75 100
B plant density
100 75 50 25 0%
Fig. 7 No competition; intercropping yield is equal with mono cropping yield and then farmer has
not relish for use of intercropping
A plant B plant
A plant density
%0 25 50 75 100
B plant density
100 75 50 25 0%
Fig. 8 Mutual inhibition; inter specific competition is higher than intra specific competition or
allelopathy or plant roots in intercropping have negative effects on the other
6.2 Mutual Inhibition (Competition)
In this case, inter specific competition is higher than intra specific competition or
allelopathy or plant roots in intercropping have negative effects on the other
(Fig. 8).
6.3 Mutual Cooperation (Protocooperation)
In this case, Intra specific competition is higher than inter specific competition. This
case is better than the other (Fig. 9).
Intercropping 135
A plant B plant
A plant density
%0 25 50 75 100
B plant density
100 75 50 25 0%
Fig. 9 Mutual cooperation; intra specific competition is higher than inter specific competition.
This case is better than the other
A plant B plant
A plant density
%0 25 50 75 100
B plant density
100 75 50 25 0%
Fig. 10 Compensation; increased amount of B plant yield is equal to decreased amount of A plant
yield in inter cropping thus A and B inter cropping yield remains unchanged
6.4 Compensation (Predation and Parasitism)
In this case, increased amount of B plant yield is equal to decreased amount of

A plant yield in inter cropping thus A and B inter cropping yield remains unchanged
(Fig. 10).
6.5 Positive Compensation
In this case, B plant yield will increase and A plant yield will decrease at inter crop-
ping system but B plant yield increase is higher than A plant yield decrease
(Fig. 11).
A plant B plant
A plant density
%0 25 50 75 100 B plant density
100 75 50 25 0%
Fig. 11 Positive compensation; B plant yield will increase and A plant yield will decrease at inter-
cropping system but B plant yield increase is higher than A plant yield decrease
A plant B plant
A plant density
%0 25 50 75 100
B plant density
100 75 50 25 0%
Fig. 12 Negative compensation; B plant yield increase and A plant yield decrease at inter cropping
system but A plant yield decrease is higher than B plant yield increase
6.6 Negative Compensation
In this case, B plant yield increase and A plant yield decrease at inter cropping sys-
tem but A plant yield decrease is higher than B plant yield increase (Fig. 12).
6.7 Unilateral Cooperation (Neamatollahi’s Definition)

(Commensalism)
In this case, A plant yield is equal in mono and inter cropping but B plant yield in
intercropping increase thus A and B inter cropping yield increase and farmer relish
for use of inter cropping (Fig. 13).
Intercropping 137
A plant B plant
A plant density
%0 25 50 75 100
B plant density
100 75 50 25 0%
Fig. 13 Unilateral cooperation (Neamatollahi’s definition); A plant yield is equal in mono and
intercropping but B plant yield in intercropping increase thus A and B intercropping yield increase
and farmer relish for use of intercropping
A plant B plant
A plant density
%0 25 50 75 100 B plant density
100 75 50 25 0%
Fig. 14 Unilateral inhibition (Neamatollahi’s definition); A plant yield is equal in mono and inter-
cropping but B plant yield in intercropping decrease thus A and B intercropping yield decrease and
farmer has not relish for use of intercropping
6.8 Unilateral Inhibition (Neamatollahi’s Definition)

(Amensalism)
In this case, A plant yield is equal in mono and inter cropping but B plant yield in
intercropping decrease thus A and B inter cropping yield decrease and farmer has
not relish for use of inter cropping (Fig. 14).
For better understand of plants relationship in intercropping we used of two
tables that first table is about the relationships between plants in intercropping and
then second table that is about the relationships between organism in world of nature
(Tables 3 and 4).
Table 3 The relationships between plants in intercropping (Neamatollahi’s definition)

Intercropping
Relations Plant A Plant B Plants A + B
No competition 0 0 0
Mutual inhibition − − −
Mutual cooperation + + +
Compensation − + 0
Positive compensation − + +
Negative compensation − + −
Unilateral cooperation (Neamatollahi’s definition) 0 + +
Unilateral inhibition (Neamatollahi’s definition) 0 − −
Table 4 The relationships Unique Society

between organisms
Relations A B A B
Mutualism − − + +
Protocooperation 0 0 + +
Commensalism − 0 + 0
Competition 0 0 − −
Amensalism 0 0 0 −
Parasitism − 0 + −
Predation − 0 + −
Noutralism 0 0 0 0
7 Conclusion
Annual intercrops have been reported to be more productive than comparable sole
crops (Hauggaard-Nielsen et al. 2001a; Hauggaard-Nielsen et al. 2001b; Jensen
1996b; Ofori and Stern 1987; Willey and Osiru 1972). Variety of opinions about the
functions of diversity in agricultural cropping systems is high while the data on
which a solid judgment could be formulated remains sparse (Giller et al. 1997;
Swift and Anderson 1993; Vandermeer et al. 1998). However it seems to be an agree
must that crop-species composition and diversity may among other things profoundly
affect soil fertility (Russell 2002; Swift and Anderson 1993), increase nutrient and
water-use efficiency and resistance to crop diseases (Mitchell et al. 2002), thereby
providing stability to the cropping system (Swift and Anderson 1993; Trenbath
1999). Farmers have generally regarded intercropping as a technique that reduces
risks in crop production as if one member of an intercrop fails, the other survives
and compensates in yield to some extent, allowing the farmer an acceptable harvest.
Pest levels are often lowered in intercrops, as the diversity of plants hampers
movement of certain pest insects and in some cases encourages beneficial insect
populations. Intercropping can be viewed from ecological, social and economic
perspectives and should be assessed relative to all three.
Intercropping 139
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Enhancing Fertilizer Efficiency in High Input
Cropping Systems in Florida
Johannes M.S. Scholberg, Lincoln Zotarelli, Michael D. Dukes,

Monica Ozores-Hampton, Guodong Liu, and Pablo Tittonell
Abstract During the last century, a number of strategies have been used to determine
optimal N-fertilizer rates and to develop appropriate N-fertilizer recommendations
for intensively-managed cropping systems. However, these strategies lack a system-
based approach and the precision needed to warrant high yields while addressing
environmental concerns in a cost-effective manner. Therefore, a more holistic
approach is required to enhance fertilizer use efficiency (FUE) in high input agricultural
systems that pose both large environmental and economic risks. This article presents
a physiological basis for improving FUE in these systems by linking physiological
crop nutrient requirements with nutrient uptake efficiencies as affected by root
characteristics, crop N demand, and production management practices. Starting at
the crop and field level we outline key processes affecting crop N demand and
uptake efficiency. For this purpose we reviewed key scientific papers that describe
yield response and fertilizer uptake efficiencies with special reference to pepper
(Capsicum annuum L.), potato (Solanum tuberosum L.) and tomato (Lycopersicon
esculentum L.) crops in Florida production systems. This because such systems are
especially prone to N leaching. Based on this review it is evident that yield response
to fertilizer for most crops tend to be inconsistent both within and across locations.
Therefore, use of standard recommendations may not be appropriate since they pose
substantial economic and environmental risks.
J.M.S. Scholberg • P. Tittonell

Wageningen University, Organic Farming Systems, Droevendaalsesteeg 1,
Wageningen, 6708 PB, The Netherlands
L. Zotarelli (*) • M. Ozores-Hampton • G. Liu
Horticultural Sciences Department, University of Florida, Fifield Hall,
Gainesville, FL 32611, USA
e-mail: lzota@ufl.edu
M.D. Dukes
Agricultural and Biological Engineering Department, University of Florida,
Frazier Rogers Hall, Gainesville, FL 32611, USA
144 J.M.S. Scholberg et al.
In terms of production efficiencies, at low fertilizer application rates values were

100–397, 63–243, 82–264 kg extra yield per kg fertilizer for tomato, pepper, and
potato, respectively. Corresponding values at recommended fertilizer rates were
reduced to 24–213, 30–152, 48–173 kg extra yield per kg fertilizer. However, using
an economic yield analysis it was shown that under adverse conditions, use of higher
fertilizer rates is correctly perceived by farmers as viable strategy for minimizing
economic risk. In terms of uptake efficiency, even at low N-fertilizer rates the fraction
of applied fertilizer that was removed by the crop was highly variable, with values
being 43–71%, 16–71%, and 4–81% for tomato, pepper, and potato, respectively.
Moreover, overall crop N recovery tended to decline with increasing N-application
rates. Since any residual soil-N may be readily lost by soil via leaching, this implies
that even at low fertilizer application rates, environmental impacts may still occur.
Furthermore this environmental risk is greatly increased at higher application rates.
These results thus are in contrast with those for the economic analysis, thus pointing
to potential stakeholder conflicts. Integration of different data sets to elucidate more
generic trends showed that changes in relative N uptake during the crop growth
cycle followed distinct patterns. These patterns were relatively consistent across
crops, years and locations. Thereby they may be used as a scientific base to structure
tactical guidelines for more efficient in-season fertilizer management based on
actual crop growth processes. This insight is especially useful for enhanced design-
ing of fertilizer applications schemes or the engineering of controlled release fertil-
izer materials. Via improved synchronization of nutrient supply with crop demand
both economic and environmental production goals thus may be attained. Such
approach thereby may afford producers with cost-effective production options
that can be readily integrated in future best management practices for high input
cropping systems.
Keywords Nitrogen uptake efficiency • Fertilizer management strategies • Intensive

management systems fertilizer recovery • Nitrogen leaching • Environmental quality
• Best management practices
1 Introduction
In the context of this article we will make special reference to pepper (Capsicum
annuum L.), potato (Solanum tuberosum L.) and tomato (Lycopersicon esculentum L.)
as examples of economically important crops. Due to their high value, these crops are
typically receiving large amounts of fertilizer rates to minimize the risk of yield reductions
due to nutrient deficiencies. Moreover, these crops are also grown within a limited
period of time, and especially during the initial part of the growing season they lack
extensive root systems. As a result their root nutrient extraction efficiencies thus may
be low and crops may not make effective use of soil nutrients. The combination of
high fertilizer application rates, low nutrient extraction efficiencies and potentially
large fluxes of irrigation water and rainfall greatly increases the risk of N leaching.
Enhancing Fertilizer Efficiency in High Input Cropping Systems in Florida 145
In this paper we also place special emphasis on Florida production systems on sandy
soils. This because these agroecosystems feature high inputs of water and nutrients
and thus are inherently prone to N leaching. Due to their low inherent soil fertility
status these systems are also uniquely suited for assessing crop fertilizer responses.
In terms of general production settings, Florida ranked first in the United States
in production value of fresh-market tomatoes, second for bell peppers and seventh
for potatoes (NASS 2012). The crop value of these three crops altogether was US
$884.9 million, which represented more than 50% of the value of Florida’s vegeta-
ble crops (FDACS 2007; NASS 2012). The statewide production area of pepper,
tomato and potato amounts to 36,340 ha. The predominant soil type in Florida
consists of coarse-textured soils with low water holding capacity and soil organic
matter content (Cantliffe et al. 2006).
Irrigation methods can be divided in two groups being low volume systems includ-
ing drip irrigation and subsurface or “seepage” irrigation. Seepage irrigation implies
maintaining a perched water table at 90–150-cm below the soil surface. This is
achieved by pumping ground water into canals or ditches spaced 25–35 m apart. Due
to the presence of an impervious soil layer at a soil depth of 1.5–2 m that effectively
hampers rapid percolation, a shallow water table thus may be maintained. (Simonne
et al. 2010). Due to Florida’s humid climate, use of drainage systems and sloped
fields are necessary for surface drainage in seep-irrigated cropping systems, which
can lead to surface runoff due to the slope. Overall, seepage irrigation has low water
use efficiency, since large volumes of water are required to maintain a shallow water
table throughout the crop season. The use of large volumes of water in turn hamper
efficient fertilizer use since nutrient losses via lateral and vertical drainage may be
substantial (Pandey et al. 2007). However, seepage irrigation is a commonly used
irrigation system for potato primarily due to its low operation costs and since it is
easy to manage. Approximately 44% of Florida irrigated area is under seepage irriga-
tion (Fig. 1). Conversely, producers aiming to enhance production efficiency in terms
use of increasingly scarce water resource may opt to use drip irrigation instead. This
irrigation system is commonly used for the production of pepper and tomato (Fig. 2).
Drip irrigation systems distribute water to a small section of the crop root zone thus
lowering water losses by wind drift and evaporation (Boman and Obreza 2002;
Locascio 2005). Additional advantages include increased flexibility of timing of fer-
tilizer application through the injection of fertilizer in the irrigation systems, precise
water distribution, and the potential use of small pumps for electronically scheduled
irrigation of large areas. Use of drip irrigation resulted in higher application
efficiencies for vegetables in Florida (80–90%) compared with seepage (20–50%)
(Simonne et al. 2010). The combination of drip irrigation and improved irrigation
scheduling techniques has been shown to greatly increase irrigation water use
efficiency and thus reduce irrigation water requirements by over 30%. With drip
irrigation small volumes of water are being applied more frequently, which can
reduce water stress and excessive percolation. Moreover, irrigation is only applied to
specific soil sections where roots tend to proliferate. Due to the synergetic effect of
these processes fertilizer is retained in the effective root zone longer and water and
nutrient resources are used more efficiently. As a result, growers can attain maximum
Fig. 1 Photograph of typical potato field under seepage irrigation in northeast Florida. The irriga-
tion is primarily the result of upward movement of water from capillarity resulting from artificial
maintained water table. This water table is maintained by water furrows where an outlet from a
pressurized source is used to deliver water to the furrow normally spaced 18–20 m and there by
maintaining a shallow water table
yields at much lower water and fertilizer application rates while potential N-leaching
losses thus minimize water quality impacts of production systems.
In terms of fertilizer use, with seepage systems, the majority of fertilizer is being
applied prior to planting with the remainder being applied as a side dressing for
potato. With the use of plastic mulched bedding systems additional fertilizer may be
applied during the season by using a fertilizer wheel, which injects solid fertilizer
by punching small holes in the plastic mulch. With the use of drip irrigation approx-
imately 20–40% of the N is applied to the soil during bed preparation. The remain-
der is commonly applied via weekly or daily injections of liquid fertilizer into the
irrigation system (fertigation).
Commonly used fertilizer materials include ammonium nitrate, ammonium sul-
phate, calcium nitrate, diammonium phosphate, potassium nitrate and urea. Use of
organic amendments including compost are typically limited.
Although there is a wealth of information on crop yield response to different fer-
tilizer rates, they fail to provide a clear understanding of the underlying crop growth,
environmental and managerial processes that control fertilizer uptake efficiency and
thus crop yield. The objective of this article is to provide a comprehensive overview
of N management approaches for peppers, potatoes and tomatoes since these crops
are similar in genetic traits, crop ecology, and production practices. The specific
Fig. 2 Photograph of traditional soil preparation of plastic mulch raised bed with drip irrigation
for tomatoes and peppers in southwest Florida. The beds are typically 15–25 cm height and
60–90 cm in width. The plastic mulch serves to protect and retain water and nutrients, minimize
weed growth, maintain bed shape
objectives of this paper are to (1) outline prevailing nutrient application philosophies;
(2) link physiological and management aspects of N-fertilizer use efficiency; (3)
compare and contrast N uptake efficiencies for different solanaceous crops; and (4)
develop functional relationships that can provide a scientific basis for improved syn-
chronization between fertilizer application and physiological N requirements; and
(5) Propose strategies for enhancing fertilizer use efficiencies in agricultural produc-
tion systems.
2 Nitrogen Use Efficiency and Crop Production
Maximizing crop yields and production efficiency have resulted in increased avail-
ability of high quality food products at a minimal cost, but public opinion may not
always reflect a positive appreciation of these achievements of modern agriculture.
During the last decades, increased public awareness of the contribution of agricul-
ture to non-point pollution has resulted in increased concerns and more stringent
environmental regulations.
Nitrogen uptake efficiency (NUE) is closely associated with non-point pollution.

For non-leguminous crops such as bell pepper, potato, and tomato, it includes two key
components. The first being efficiency of roots to take up plant available N from the
soil. The second is the conversion efficiency. This is a measure of how effective crops
are in converting N taken up by the roots into overall plant material or harvestable
products. In terms of overall fertilizer use, the term fertilizer N recovery which is the
fraction of applied N fertilizer take up by the crop is especially pertinent. This since it
is a measure of resource use efficiency as related to N-fertilizer being used. Moreover,
crop N recovery tends to be also inversely linked to both residual soil N and N emission
losses. The last being a proxy of potentially negative environmental impacts.
Nitrogen uptake rate of roots depends on three genetic characteristics (Tinker and
Nye 2000). The first being maximum uptake rate or Vmax. This is the maximum root
N-uptake rate at very high nutrient concentrations and is indicative of uptake
capacity. High Vmax values imply that roots can make effective use of soil nutrients at
high soil concentrations. This since roots take up nutrients quickly before they may
be lost due to leaching. The second trait is extraction efficiency which is referred to
as Km. This is the nutrient concentration at which actual uptake is half the potential
uptake rate. Low Km values imply that plants can still sustain relatively high nutrient
uptake at low soil solution concentrations. Lastly, the compensation point or Cmin is
the concentration below which plants no longer effectively extract nutrients from
the soil solution. So very low Cmin values imply that crops are excellent nutrients
scavengers. The last two traits thus are indicative of the inherent potential nutrient
uptake efficiency of a crop. Although these traits are mainly determined by inherent
genetic characteristics, actual N-uptake rate also depend on crop water use, crop N
status and the availability of assimilates as well (Scholberg and Morgan 2012).
In addition to genetic traits, irrigation and fertilization management play a major
role in terms of determining solute concentrations and movement of water and
nutrients across the soil-plant-atmosphere continuum. Thereby, irrigation or fertil-
ization method and timing can contribute significantly to overall N recovery. In
terms of N conversion efficiency this consists of two key components. The first
being N transfer and re-translocation which includes remobilization of proteins
from existing tissue with reduced metabolic activity including senescing leaves and
storage organs. The second pertains to carbon fixation rates per unit of taken up N.
This is affected by genetic traits including metabolic pathways, inherent cellular
structures, tissue functionality, metabolic activity but also environmental conditions
such as crop light, water and nutrient status. In this context both irrigation and
fertilization management can greatly affect both carbon and N assimilation cycles
through various feedback mechanisms. Since both water and nutrients commonly
are the most limiting production factors, sound irrigation and fertilization manage-
ment is required in order to ensure maximum crop yields and profitability of com-
mercial crop production systems. Therefore, appropriate production guidelines are
needed to assist crop producers to increase the efficiency of use of N-fertilizers
especially in high value cropping systems. In practice, economic N conversion
efficiency and/or N recovery may be more relevant for crop producers or policy-
makers. This since it expresses N utilization on a marketable commodity basis rather
than on a total crop biomass basis. In this case efficiency is being defined either
as incremental marketable yield produced per unit of applied N fertilizer or as
incremental yield per unit extra N accumulated by the crop. Overall nitrogen use
efficiency thus is affected by inherent genetic traits, environmental conditions and
management practices. Since water and nutrient efficiencies are intrinsically linked,
both irrigation and fertilizer managements need to be optimal to warrant optimal
yields and maximum resource use efficiencies.
3 Overview of Prevailing Nutrient Application Philosophies
Historically, soil fertility research has focused on yield response to fertilizer rates in
order to maximize production and/or net returns for selected cropping systems and
specific pedoclimatic conditions. These approaches have been used to assess how
yields are affected by fertilizer rates. Simple statistical analysis may be then used to
infer which fertilizer rate may be optimal. This is exemplified using existing data
sets from the literature (Locascio et al. 1996; Scholberg et al. 2000a) as is shown in
Fig. 3. In this manner we can generate basic fertilizer response curves and by using
a quadratic model, maximum yields can be estimated. This is done by setting the
first derivative of the quadratic equation generated by the regression analysis equal
to zero. In our example resulting in Nyield optimum = 279 and 253 kg N ha−1 for both
studies, respectively. A more conservative approach is the use of a linear-plateau
method which typically results in lower N recommendations. In our specific example
the optimum N-fertilizer rate would be 165 kg N ha−1. So in both cases we use
statistical tools to infer if numeric differences among different observations are
related to a specific fertilizer effect or more likely attributed to random variation
which is a standard practice in fertilizer rate studies. So although growers may argue
that actual crop yield and profits increased up to 267 kg N ha−1 (Table 1), researchers
tend to look for scientific evidence based on standard statistics. Scientists thus
would argue that applying N-fertilizer in excess of 202–269 kg N ha−1 will not
result in significant tomato yield response. Although results may not be statistically
different, economically they may still benefit growers even if the outcome would
only occur 1 out of 5 growing seasons. So in this context, the stakeholders’ perspective
thus may bias the interpretation of the research findings.
A second consideration for assessing optimal fertilizer rates would be to focus on
economic profits. For tomato production in Florida in 2007, the average fertilizer
cost was $1.17 kg−1 of N (USDA 2010) compared to the average price of $932 Mt−1
of fresh tomato (NASS 2010). Combining this information with the yield function
(Scholberg et al. 2000a, b) shown in Fig. 3 and by multiplying the yield values by
the unit crop price a crop value (V) function can be developed:
V = 20,519 + 339 N - 0.6761 N 2

Where “N” is the amount of fertilizer being applied in kg N per ha.
Tomato yield
(Mt ha-1)
80
Scholberg et al. (2000)
60
Linear Plateau (LP)
Locascio et al. (1996)

40
20
0
0 50 100 150 200 250 300 350
N-fertilizer rate (kg N ha-1)
Fig. 3 Effects of N-fertilizer rate on marketable fruit yield and optimal N-fertilizer rates using qua-
dratic (QM) and linear-plateau (LP) approach (Adapted from Scholberg et al. 2000a, b and Locascio
et al. 1996). The figure is an example of the use of simple statistical analysis to infer which N-fertilizer
rate may be optimum, basic fertilizer response curves can be generate by using a quadratic model,
thus maximum yields can be estimated. By setting the first derivative of the quadratic equation gener-
ated by the regression analysis equal to zero. Locascio et al. (1996): QM: Y = 14 + 0.33x − 0.0006x2,
Nopt = 279 kg N ha−1; Scholberg et al. (2000a, b): QM: Y = 22 + 0.36x − 0.0007x2 Nopt = 253 kg N ha−1;
LP:Y = 13 + 0.29 N (N < 165) Y = 61 (N > 165), Nopt = 165 kg N ha−1
Table 1 Effect of nitrogen (N) rate on tomato yield, net profit, marginal N efficiency (increased
yield/unit N fertilizer) and residual soil N from fertilizer. With an increase in N-rate from 133 to
333 kg N ha−1, the marginal crop N efficiency, which is the yield increment per unit extra fertilizer
being added, dropped from 265 to 30 kg of tomato per kg N. At a fertilizer rate of 133 kg N ha−1
the amount of N not taken up by the crop was 49 kg N ha−1 but this amount increased to 207 kg N
ha−1 for the highest N-rate. This underlines that maximizing production by increasing the N-rate
will invariably result in less efficient fertilizer use; with a decrease in fertilizer use efficiency an
increasingly large fraction of fertilizer N can be potentially lost from the production system
Fertilizer rate Tomato yield Net profita Marginal N-efficiency Residual soil N
(kg N ha−1) (Mt ha−1) ($ha−1) (kg Y kg N−1) (kg N ha−1)
133 57.8 26,337 265 49
200 66.2 33,798 124 107
267 68.2 35,317 77 157
333 63.9 30,896 30 207
a
Assuming a production cost of $27,244 ha−1 and tomato prices of $932 Mt−1
The fertilizer cost function can be defined as follows:
C = a + 1.17 N
With “a” being application cost and “N” being the amount of fertilizer being
applied in kg N per ha. Profits are maximized if marginal value by equating mar-
ginal value (dV/dN) to marginal cost marginal cost (dC/dN). This translates to:
339 - 2 * 0.6761 N = 1.17
Solving for N results in Necon.optimum = 250 kg N ha−1. Since fertilizer costs make up a
relatively small fraction (3.5%) of total production cost for tomatoes or peppers,
economic and yield optima thus appear to be nearly identical. A recent yield and
economic assessment for processing tomatoes in Canada resulted an economic
optimum N rate values of 5–8 kg N ha−1 lower than the N-rate of 271 kg N ha−1
required for the maximum marketable yield (Zhang et al. 2010).
A third consideration is including the highly stochastic nature of environmental
conditions and crop management and their effects on N uptake dynamics, crop
growth and yield. Each time a fertilizer rate study is conducted a specific optimal
N-rate can be determined that is relevant for those set of conditions. However, this
optimum may greatly vary from year to year or even within a single field. A literature
review of fertilizer response curves on tomatoes for the same location (for example
Quincy, Florida), showed optimal N rates to be 125 and 226 kg N ha−1 in two
consecutive years (1983 and 1984) with respective maximum yields of 88 and
81 Mt ha−1 (Scholberg et al. 2000a, b). An effect of adverse environmental condi-
tions on tomato production could be observed in a 3-year experiment conducted
in Citra, Florida, USA between 2005 and 2007 with drip irrigated/fertigated tomato.
No statistical differences in marketable tomato yield among N-rates of 176, 220 and
330 kg N ha−1 within experimental years were reported. Even though crop management
and irrigation were consistent across years, average marketable yield was 28, 56 and
79 Mt ha−1 in 2005, 2006 and 2007, respectively. So it is evident that non-fertilizer
related factors including appreciable differences in temperature, rainfall and solar
radiation played an important role. This in turn drastically affected plant growth and
development and determined overall fruit yield potential (Zotarelli et al. 2009b).
Based on N rates to produced optimal yield and low cost of N-fertilizer, growers may
opt to use excessively high fertilizer rates as an “inexpensive insurance policy” to mini-
mize the risk of yield losses due to N limitations under adverse conditions. Using results
from a survey of reported yield response curves to N rates for drip irrigation based sys-
tems for five locations with similar production systems in the southeastern U.S.
(Scholberg et al. 2000a, b) a mean for optimal N rates of 221 kg N ha−1 with a standard
deviation of 64 kg N ha−1was obtained. Assuming that 95% of possible outcomes occur
within two standard deviations from the mean, the following would hold:
93kgN ha -1 < N yield optimum < 349kgN ha -1

Using the yield equation presented in Fig. 3 the extreme cases would be:
Y(93N) = 22 + 0.36 * (93) - 0.0007 * (93)2 = 49.4Mt ha -1 versus
Y(349N) = 22 + 0.36 * (349) - 0.0007 * (349)2 = 62.4Mt ha -1
Assuming an overall production cost of $27,244 ha−1(Olson et al. 2010) and

average crop value of $932 Mt−1 of tomato (NASS 2010) net profits would be:
(49.4 * $932) - $27,244 = $19,262 ha -1 versus

(62.4 * $932)- $27,244 = $30,913 ha -1
for the “low” versus “high” N-fertilizer rate scenario, respectively. The additional
economic cost associated with a more risk-adverse approach would be:
(349 - 93) * $1.17 = $299
However, this may be resulting in a net potential profit gain of $11,651, which
underlines that over-fertilization is correctly perceived as a relatively cost-effective
insurance premium by growers. However, when the economic approach is consid-
ered as strategy for N-fertilization management, with the Necon. optimum = 249 kg N ha−1,
net profits would be as follows:
(67.6 * $932) - $27,244 = $35,759 ha -1 ,
Thus resulting in $4,846 ha−1 more or 13.5% higher profit compared to a N-rate of
349 kg N ha−1. Based on this the optimal rate under standard conditions would be
249 kg N ha−1 but growers may still opt to apply more based on their perception
and risk aversion. However, growers currently are also increasingly held liable for
externalities associated with their operations. This in turn adds complexity to the
overall decision-making process.
For the same example crop discussed above, N uptake and fertilizer use efficiency
were assessed. With an increase in N-rate from 133 to 333 kg N ha−1, the apparent N
recovery, which is an estimate of fertilizer uptake efficiency, dropped from 0.65 to
0.39 (Fig. 4). Total crop N uptake from fertilizer increased from 85 to 129 kg N ha−1.
The marginal crop N efficiency, which is the yield increment per unit extra fertilizer
being added, dropped from 129 to 11 kg of tomato per kg N as N-fertilizer rates
increased from 133 to 333 kg ha−1 (Table 1). This underlines that maximizing pro-
duction by increasing the N-rate will invariably result in less efficient fertilizer use
(Zotarelli et al. 2009a, b). With a decrease in fertilizer use efficiency an increasingly
large fraction of fertilizer N can be potentially lost from the production system. At
a fertilizer rate of 133 kg N ha−1the amount of N not taken up by the crop was 49 kg
N ha−1 but this amount increased to 207 kg N ha−1 for the highest N-rate (Table 1).
There are only two multiple N-rate studies with seepage irrigation conducted in
Florida by Hochmuth et al. (1989) and Everett (1976). The limitation of one of the
N uptake or loss (kg N ha-1)

250 0.8
ARN
N potential loss 0.7
Apparent N recovery (ARN)

200
0.6
150
0.5
0.4
100
N uptake from
fertilizer 0.3
50
0.2
0 0.1
0 50 100 150 200 250 300 350
N-fertilizer rate (kg N ha-1)
Fig. 4 Effects of N-fertilizer rate on tomato N uptake from fertilizer, potential N loss/leaching
from fertilizer, and apparent fertilizer uptake efficiency (Adapted from Scholberg 1996 and
Locascio et al. 1996). Nuptake = 2 + 0.68x − 0.001x2, r2 = 0.98; Nloss = − 54 + 0.78x, r2 = 0.99. The graph
shows that maximizing production by increasing the N-rate will invariably result in less efficient
fertilizer use. With a decrease in fertilizer use efficiency an increasingly large fraction of fertilizer
N can be potentially lost from the production system
studies was the narrow N rate considered from 180 to 450 kg. ha−1. In addition to this
the tomato yields in both studies were well below current production levels. Therefore,
a study was designed to identify a range of N rates that would result in optimal yields,
acceptable postharvest quality, and maximum economical return for spring-grown
tomato with the use of seepage irrigation (Ozores-Hampton et al. 2012). In this
study in Palmetto, Florida, USA, N rates were ranging from 22 to 470 kg· ha−1.
Fertilizer was applied pre-plant as ammonium nitrate. While interpreting results
absence of heavy rainfall events and frost protection was important since both will
result in raised water table levels and a significant loss of fertilizer N. Provided such
conditions did not prevail, extra-large and total marketable fruits yields showed a
quadratic plateau response to N rates maximum yields occurred between 172 and
298 kg N· ha−1. Post-harvest fruit quality measured as fruit soluble solids concen-
tration, total titratable acidity and pH at harvest was not affected by N rates. Values
ranged between 5.53° and 3.63° Brix, 0.57 to 0.31 mEq/100 g juice, 4.61–4.43, for
2007 and 2008, respectively. In 2007, the N-rate required for maximum total
marketable yield was 172 kg ha−1. This value is 23% lower than the current recom-
mended rate of 224 kg ha−1. Given the functional form of the yield response curve
(quadratic-plateau), applying an additional 52 kg ha−1 of N fertilizer did not result in
significantly higher tomato yields. In 2008, growing conditions were less favorable
Fig. 5 Tomato (‘Florida 47’) yield response to N rates during the spring of 2007 (a) and 2008 (b)
for total marketable (three harvests combined), following a quadratic-plateau model: Y = a + bX + cX2
if x < N critical rate, y = Plateau yield if X > N critical rate. Each data point is a mean of four replica-
tions consisted of marketable mature-green and colored tomatoes graded in the field according to
USDA specifications. Bars denote standard error. (Adapted from Ozores-Hampton et al. 2012). (a)
2007: Ytotal = 35.7067 + 0.7538 X − 0.0022 X2, r2 = 0.74; Ymkt = 21.3359 + 0.8764 X − 0.00322 X2;
r2 = 0.63; (b) 2008: Ytotal = 45.8748 + 0.2713X − 0.00045 X2; r2 = 0.78; Ymkt = 38.5395 + 0.1833X −
0.00033 X2; r2 = 0.67
and the calculated optimal N-rate to maximize marketable tomato yields was
298 kg ha−1. This value is 74 kg ha−1 above of the current recommendation (Fig. 5).
The addition of this extra fertilizer was estimated to increase maximum tomato yields
by more than 2.5 Mg compared to the use of the recommended rate (Fig. 5). Therefore,
from an economic perspective using excessively high fertilizer rates may pay off during
adverse conditions. The extremely high price ratio of fresh market tomatoes relative
to the fertilizer-N cost appears to provide a clear incentive to growers to apply nearly
50% more N compared to the recommend N rate.
In terms of environmental economics, Florida sandy soils have low water and nutrient
holding capacities. Therefore, a substantial fraction of this unused fertilizer may
ultimately end up in the groundwater (Huang et al. 1994). It is thus anticipated that
fertilizer-N uptake efficiency will become a critical consideration in the overall assess-
ment whether vegetable production will be a viable and sustainable land use option on
vulnerable soils that are prone to N leaching. It is apparent that yield response to
fertilizer application rates for most crops tend to be inconsistent both within and across
locations. Therefore, use of standard recommendations may not be appropriate since
they pose substantial economic and environmental risks. However, growers do perceive
the risks of adverse conditions reducing their crop yields to be a significant threat. This
in combination with the very high tomato price to fertilizer cost ratio may entice them
to be using excessively high fertilizer rates as affordable strategies to prevent yield losses
under unfavorable production conditions.
4 Linking Crop Physiology and Production Efficiency
Nitrogen is a key nutrient required to maximize yields of high value crops. Most
vegetable crops have relative low nutrient uptake rate during the initial growth
period followed by an exponential increase in nutrient requirements towards the
crop maturity(Huett and White 1992). During crop maturation stage, nutrient uptake
rate is slow again and N uptake thus follows a sigmoid curve. This is particularly
true of the N requirements of tomato, potato and bell pepper. In terms of their cor-
responding nutrient extraction efficiencies, approximately 80–90% of their roots are
located in the upper 15–30 cm of the soil profile(Scholberg 1996; Munoz-Arboleda
et al. 2006; Zotarelli et al. 2009a, b). Once N moves below the root zone it may
become a liability rather than an essential nutrient used to increase crop yield. For
soil types vulnerable to N leaching, improved irrigation and fertilizer management
practices are thus essential to ensure maximum fertilizer uptake. This in turn is cru-
cial to minimize environmental impacts of commercial vegetable production opera-
tions. But this requires a better understanding of the processes that control crop
fertilizer uptake and how this relates to overall N-fertilizer recommendations.
Nitrogen recommendations integrate two intrinsically linked aspects. The first
one is crop physiological N requirement or the overall crop N content at which N is
no longer limiting growth or yield. This value is defined by inherent genetic traits
and plant growth conditions that in turn set the overall yield potential. The second
component is the actual fertilizer uptake efficiency for a specific production system.
This is defined by environmental conditions and management practices. Combining
these components results in a general N-fertilizer recommendation as follows:
N recommendation = crop N requirement / N-fertilizer uptake efficiency.

The first component can be determined from overall crop N accumulation under
optimal growth conditions. Values may vary depending on crop vigor as affected by
genetic traits and (a)biotic stress factors. The second component is rather complex
in nature since it includes crop type, soil conditions, plant age, irrigation management,
fertilizer management and environmental factors (e.g. radiation and temperature).
Due to the dynamic and interactive nature of processes that control uptake
efficiencies, it is difficult to ascertain its value. On the other hand, it is quite important
since it is inversely related to potential N loading rates of water resources. In summary
by using our conceptual framework it is evident that crop N recommendations
embody the interaction between crop physiological N demand and uptake efficiency.
In this manner we aim to make the linkage between production goals, genetic pro-
duction potential, crop N accumulation, and resource management more transparent.
The underlying mechanisms and processes involved will be discussed in the more
detail in subsequent sections.
5 Crop N Requirements
5.1 Overall N Accumulation
Crop N demand is the product of dry matter accumulation of plant components and
their respective N concentration. Therefore, the extent of carbon accumulation and
the N content of dominant plant tissues have a major role in setting the total N
uptake capacity for a specific crop. This demand is primarily driven by shoot
growth since shoots form the greatest N-sink due to their high N content (Thorup-
Kristensen and Sorensen 1999). Root dry matter constitutes a minor part of the
total cumulative output of dry matter in potato (excluding tuber weight), tomato
and bell pepper crops (Lesczynski and Tanner 1976; Jackson and Bloom 1990;
Olsen et al. 1993; Tapia and Gutierrez 1997; Vavrina et al. 1998). For these crops
there is a linear relationship between cumulative shoot dry matter and N accumula-
tion (Miller et al. 1979; Millard et al. 1989; Tapia and Gutierrez 1997; Belanger
et al. 2001; Alva et al. 2002).
While N uptake of these crops is initially set by the magnitude of their vegetative
shoot growth, this demand can exceed the necessary requirement to produce the
optimum yield of fruit or tubers. Excessively high N-rates reduce the N uptake
efficiency by which plants convert fertilizer N into plant constituents (Millard and
Marshall 1986). Carbon cost associated with N uptake and storage may actually
reduce overall productivity and/or partitioning of assimilates to fruits and tubers
(Kleinkopf et al. 1981; Westermann and Kleinkopf 1985; Millard and Marshall 1986;
Scholberg et al. 2000a, b; Cambouris et al. 2008). Maximum reported N accumula-
tion rates were on the order of 270–350 kg N ha−1 for potato (Kleinkopf et al. 1981;
Roberts et al. 1991; Alva et al. 2002), 193–234 kg N ha−1 for pepper (Locascio et al.
1985; Tei et al. 1999), and 159–350 kg N ha−1 for tomato(Baryosef et al. 1980;
Andersen et al. 1999; Tei et al. 2002; Hartz and Bottoms 2009). However, oversupply
of N can alter the normal N demand and keep over-vegetative growth during ontogeny.
This abnormality of N fertilization may hamper reproductive growth and optimal
development of economic plant parts. Furthermore, over N fertilization basically
relates disease and pest problems and thus overall production of these crops. In
potato, this problem is well-documented and it was shown that excessive early
applications can delay tuber formation and growth (Westermann and Kleinkopf 1985)
and late applications can inhibit senescence (Millard and Mackerron 1986; Millard
and Marshall 1986). In tomato, excess N application promotes vegetative growth and
delays fruit development and maturity (Scholberg 1996). With an increase in N
application rate, overall crop accumulation increases but fertilizer recovery and use
efficiency decreases(Vos 1999; Cambouris et al. 2008; Zotarelli et al. 2009a, b;
Zotarelli et al. 2011) (Fig. 4, Tables 2). It is thus may be concluded that crop growth
and yield are intrinsically linked to fertilizer N supply. Increased fertilizer applica-
tion will enhance crop growth and yield where as optimal crop growth will enhance
crop nutrient demand. However, excessive N application may delay harvest and also
potentially reduce resource use efficiency, yields and crop quality.
5.2 Crop N Accumulation Patterns
The combined demand of N for vegetative and reproductive growth has been quantified
in studies of potato (Millard and Marshall 1986; Duchenne et al. 1997; Alva et al.
2002), tomato (Baryosef et al. 1980; Scholberg et al. 2000a, b; Zotarelli et al. 2009a,
b), and bell pepper (Miller et al. 1979; Locascio et al. 1985; Santiago and Goyal 1985;
Zotarelli et al. 2011). Overall N uptake of planted potato, transplanted tomato
and pepper typically closely follows biomass accumulation and shows a lag phase
(28–42 days) when leaf area, leaf light interception and root uptake capacity are limit-
ing productivity and N uptake, followed by a linear exponential (rapid) growth phase,
which culminates at 70–98 days. After this, canopy N levels typically decline and re-
translocation of both energy and N to tubers or fruit predominates. Crop N accumula-
tion thus follows a logistic growth curve. To maximize N uptake efficiency, N supply
needs to be synchronized with the crop N requirement.
A secondary demand for N by tomato, bell pepper and potato crops is initiated at
the onset of their reproductive growth. In potatoes, the initial exponential phase of
dry weight and total demand for N in tubers occurred at approximately 55 days after
planting (Kleinkopf et al. 1981; Alva et al. 2002). There is limited information in
this area regarding new varieties of bell pepper and tomato. Exponential N demand
of bell pepper fruit commenced between 28 (Miller et al. 1979) and 63 days from
transplanting (Locascio et al. 1985). Tapia and Gutierrez (1997) found that the
exponential growth and increased N demands of tomato fruit occurs at 68 days from
emergence, which translates to 33–40 days after transplanting.
However, in most of these studies, no functional relationships between plant N
accumulation as a function of time were derived. Therefore there is a lack of generic
uptake curves for vegetable crops. Using data reported in the literature for constructing
Table 2 Effects of N-rate on tomato, pepper and potato yield, total nitrogen uptake, nitrogen accumulation in fruit, percentage N partitioned to fruit, fertilizer
158
recovery, nitrogen efficiency (yield per unit fertilizer N), and agronomic efficiency (incremental yield per unit fertilizer in comparison to control treatment)
N rate Yield Total. N Marketable N Mkt. N N-rec. N effic. Ag. effic. Comments
kg N ha−1 Mt ha−1 kg N ha−1 kg N ha−1 % % kg kg N−1 kg kg N−1
Tomato
USA, Florida, Quincy 1996 (Andersen et al. 1999)
0 30 43 25 58 n.a. n.a. n.a.
67 56 85 50 58 62 842 397 All trtm. mulched
134 65 110 58 52 50 488 266 All N applied
202 75 143 89 62 49 371 223 preplant
269 63 138 71 52 35 235 124 broadcast
(1 applic.)
USA, Florida, Quincy 1996 (Andersen et al. 1999)
0 49 96 58 60 n.a. n.a. n.a.
67 56 125 70 56 43 836 100 All trtm. mulched
134 59 152 77 51 42 439 71 All N applied
202 54 156 79 50 30 268 24 preplant
269 56 159 86 54 23 207 24 broadcast (1 applic.)
USA, Florida, Gainesville, 1996 (Locascio et al. 1996)
0 13 30 17 57 n.a. n.a. n.a. All trtm. mulched
133 51 125 59 47 71 386 286 20% N applied
200 56 123 63 51 47 280 213 pre-plant and rest
267 62 140 74 53 41 233 183 applied as fertigation
333 64 150 90 60 36 191 151 (12 applic.)
Pepper
USA, Florida, Gainesville, 2007 (Zotarelli et al. 2011)
176 24 56 25 45 34 141 n.a. Fertigation (13 applic.)
220 26 70 29 41 32 118 n.a.
330 28 78 32 41 23 84 n.a.
J.M.S. Scholberg et al.
224 38 234 60 26 36 170 n.a. Mulched (1 applic.)
224 34 108 50 46 30 150 n.a. no mulch (1 applic.)
224 25 120 52 43 42 112 n.a. Mulched (1 applic.)
Australia, Queensland, Spring 1990 (Olsen et al. 1993)
0 8 91 n.a. n.a. n.a. n.a. n.a.
70 25 102 n.a. n.a. 16 357 243 Fertigation (10 applic.)
140 37 133 n.a. n.a. 30 264 207
210 40 157 n.a. n.a. 32 190 152
280 43 193 76 39 37 154 125
Australia, Queensland, Fall 1991 (Olsen et al. 1993)
0 15 35 n.a. n.a.
70 27 85 n.a. n.a. 71 386 171 Fertigation (10 applic)
140 30 131 n.a. n.a. 68 214 107
210 35 161 n.a. n.a. 60 167 95
280 37 186 120 65 54 132 79
Italy, Perugia 1992 (Tei et al. 1999)
0 27 129 25 20 n.a. n.a. n.a.
100 34 168 50 30 39 337 63 Broadcast (3 applic.)
Enhancing Fertilizer Efficiency in High Input Cropping Systems in Florida
200 38 194 58 30 33 191 54

300 37 216 89 41 29 122 30
400 38 226 71 32 24 95 27
(continued)
159
160
Table 2 (continued)
Potato
USA, Michigan, East lancing, 1989 (Joern and Vitosh 1995a, b)
0 20 109 52 47 n.a. n.a. n.a.
56 29 111 75 68 4 510 153 3 applications
112 37 163 103 63 48 332 153 1 application
112 36 149 103 69 35 323 145 2 applications
112 35 157 93 59 43 315 136 4 applications
168 36 200 121 60 54 215 96 3 applications
USA, Michigan, Entrican, 1989 (Joern and Vitosh 1995a, b)
0 24 47 47 100 n.a. n.a. n.a.
56 39 59 63 100 21 697 264 3 applications
112 44 90 86 96 38 391 174 1 application
112 51 86 84 98 35 455 238 2 applications
112 42 86 80 93 35 378 162 4 applications
168 53 109 105 96 37 317 173 3 applications
USA, Washington, Paterson, 1982 (Roberts et al. 1991)
336 74 272 240 61 45 220 n.a.
UK, Scotland, Wartle, 1983 (Millard and Marshall 1986)
0 32 77 56 73 n.a. n.a. n.a.
50 36 97 76 78 39 716 82
150 46 152 108 71 49 305 93
J.M.S. Scholberg et al.
250 44 174 124 71 39 174 48

UK, Scotland, Rathienorman, 1984 (Millard and Marshall 1986)
0 37 68 55 81 n.a. n.a. n.a.
50 47 110 95 87 84 936 198
150 56 163 137 84 63 375 129
250 55 190 152 80 49 219 71
Germany, Bavaria, Scheyern 1996 (Maidl et al. 2002)
150 54 147 110 75 36 361 n.a. Broadcast (1 applic.)
150 57 168 128 76 47 381 n.a. Banded (1 applic.)
150 52 156 116 74 45 346 n.a. Banded (3 applic.)
Germany, Bavaria, Scheyern 1997 (Maidl et al. 2002)
150 62 225 171 76 66 413 n.a. Banded (1 applic.)
150 65 225 191 85 69 431 n.a. Banded (3 applic.)
Enhancing Fertilizer Efficiency in High Input Cropping Systems in Florida
161
N accumulation curves for potato, it appeared that overall uptake patterns were
similar except for differences in duration of the growth season. However, deviations
from general patterns both in terms of magnitude of total N uptake and seasonal
dynamics may occur. This may be related to genetic traits, extreme temperatures,
and possibly N and/or water limitations, which especially affect the overall magni-
tude of N uptake. Using a relative N accumulation which is defined as:
N rel = 100%* N accumulation / N maximum accumulation
Using this approach allowed development of more generic N accumulation curve

for potato, pepper and tomato that are no longer affected by the magnitude of overall
N accumulation (Fig. 6). In many cases there were pronounced seasonal and site
specific influences that governed variation in N uptake, N recovery or partitioning
to fruits or tubers (Table 2). However, seasonal effects on N uptake or partitioning
into harvest organs were considerably smaller when compared to the effects on total
shoot N recovery. This indicates that N uptake or partitioning into harvested organs
is mainly determined by genetic traits and thus relatively stable compared to total N
accumulation, which is also greatly affected by management and environmental
conditions (Olsen et al. 1993; Joern and Vitosh 1995a).
In terms of regional differences, in the case of potato (Fig. 6c) results were
obtained for the northeast USA, France, and Scotland. Therefore climatic variability,
and use of different varieties may have resulted in different N accumulation patterns.
For crops grown in similar climatic regions, seasonal differences appear to be
minimal (Fig. 6a, b). In case that N recommendations need to be developed for
regions with pronounced regional and seasonal climatic variation in terms of daily
temperatures, use of degree day-based uptake curves may be used instead (Scholberg
et al. 2002). In this manner generic uptake curves may be developed that are not
greatly affected by climatic conditions.
With regard to crop N partitioning, the N demand of either tubers or fruit for
these three crops typically exceeds the demand from their vegetative growth at
harvesting time (Table 2). For tomato approximately 50–60% of crop N was parti-
tioned to the fruit (Table 2). For pepper, between 30 and 65% of the N was accumu-
lated in the fruit with an average value of 39% (Table 2). For potato this range was
59–100% with an average value of 78%. The proportion of total N accumulated by
the crop partitioned to marketable plant components is highest in potato and least in
bell pepper. In many cases, N partitioning to reproductive organs occurred with
increasing rates of N fertilizer for these crops (Locascio et al. 1985; Millard and
Marshall 1986; Andersen et al. 1999). Especially under Florida conditions, soils
typically are low in organic matter (<0.5–1.0%) and residual soil N values typically
are on the order of <10 mg kg−1 (Scholberg 1996). These conditions thus are ideally
suited for developing fertilizer response curves since residual soil N content and
inherent soil fertility are very low. On the other hand, if an appreciable fraction of
crop N is left with crop residues in the field, most of this N may still be lost due to
leaching during summer fallows.
Nitrogen partitioning to marketable crop components can also be assessed by
calculating nitrogen use efficiencies. In this case, efficiencies are based on crop
yield rather than on overall biomass accumulation. Values are typically calculated
Relative N-accumulation %
a 100
Tomato
80
60
40
20
0
0 14 28 42 56 70 84 98 112
b 100
Pepper
80
60
40
20
0
0 14 28 42 56 70 84 98 112
c 100
Potato
80
60
40
20
0
0 14 28 42 56 70 84 98 112 126
Days after planting
Fig. 6 Developing a generic N accumulation curve for (a) tomato for subsurface (sub) and drip-
irrigated (drip) tomato derived from Scholberg (1996); (b) pepper from Santiago and Goyal (1985)
and Miller et al. (1979) and (c) potato derived from Alva et al. (2002); Duchenne et al. (1997); Millard
and Marshall (1986); and Locascio et al. (1996). Despite data being obtained from different studies and
production conditions, the homogeneity of the functional relationship for most crops is noticeable,
indicating that there are generic uptake trends that hold across crops and production environments.
Based on crop N accumulation patterns, tomato has a relatively short period of high uptake rates
compared to peppers and potatoes. It is proposed that on soils with limited nutrient retention and/
or with the use of hydroponics, matching fertilizer supply with crop N requirements may facilitate
more efficient N uptake efficiencies. By applying excessive amounts of pre-plant broadcast applied
fertilizer will result in high residual soil N during the initial 5–6 weeks increasing potential N-leaching
as the ratio of total yield and the amount of N-fertilizer being applied. The reason
for using this approach was that it allows a general assessment of efficiencies across
all studies also for those were only a single N rate was used. Average NUE for
representative N rates (150–250 kg N ha−1 at which yields were within 90% of
maximum yield) were 291, 166, 229 kg yield produced per kg N applied for tomato,
peppers, and potato, respectively (Table 2). Use of agronomic efficiencies (AE)
allows assessment of the overall efficiency by which N-fertilizer is converted into
marketable yield since it corrects for residual and/or mineralized soil N (Olsen et al.
1993). By using this approach, efficiencies were 179, 39, and 97 kg yield produced
per kg N applied for tomato, peppers, and potato, respectively (Table 2).
Alternatively the average fruit N harvest indices may be used which can be
defined as follows:
NHI = total N fruit/total N above-ground at harvest
This index may be greatly affected by environmental condition. In three consecutive
years, Zotarelli et al. (2009a, b) reported an overall tomato NHI of 0.36; 0.62; and 0.74
for seasons of 2005, 2006, and 2007, respectively. Lower values during 2005 were
related to unfavorable growth conditions hampering initial fruit set and canopy devel-
opment while more vigorous growth during 2006 and 2007 also resulted in enhanced
initial canopy development, greater water use and fruit yield. During the spring of
2005, hot and humid weather prevailed. The tomato reproductive period during spring
of 2005 was wetter and hotter compared to the subsequent seasons which resulted in
premature blooming and flower abortion (Rao et al. 1992; Peet et al. 1997), conse-
quently leading to lower yields. In conclusion it appears that despite potentially large
differences in total N accumulation, the general shape of relative uptake curves is rather
similar. Seasonal N uptake patterns may be affected by environmental conditions and
prevailing management practices. Overall N partitioning within the crops tends to be
mainly governed by genetic traits, provided no extreme growth stress occurs.
5.3 Actual Crop N Uptake Rates
Using the first derivative (dN/dt) of the cumulative N uptake curves, daily N uptake
rates (kg N day−1) or relative N uptake rates (% of total N day−1) can be calculated
as is shown in Fig. 7. Maximum calculated N uptake rates for potato occurred
between 10 and 12 weeks after planting and actual N uptake values ranged from
4 to 10 kg N ha day−1 (Millard and Marshall 1986; Alva et al. 2002). For pepper
maximum uptake also occurred around 10–12 weeks with maximum uptake rates
of 2.5–5 kg N ha day−1 (Miller et al. 1979; Locascio et al. 1985). Uptake rates for
tomato peaked between 8 and 10 weeks with maximum values of 5–8.5 kg N ha
day−1 (Baryosef and Sagiv 1982; Scholberg 1996; Zotarelli et al. 2009a, b). In terms
effects of soil N solution concentrations, it was reported that the maximum uptake
for tomato occurred at solution concentrations above 200 ppm NO3 (Baryosef and
Sagiv 1982).
Relative daily uptake %

3.5
Tomato
3.0
Potato
2.5
2.0
1.5
Pepper
1.0
0.5
0.0
0 14 28 42 56 70 84 98 112 126
Days after planting
Fig. 7 Comparing N uptake dynamics between potato, pepper, and tomato by using the first deriv-
ative (dN/dt) of the cumulative N uptake curves, daily N uptake rates (kg N day−1) or relative N
uptake rates (% of total N day−1) can be calculated. Maximum calculated N uptake rates for pepper
and potato occurred between 10 and 12 weeks, and uptake rates for tomato peaked between 8 and
10 weeks. It is evident that crop nutrient requirements during the initial period are very low for all
three crops during the first 6 weeks after transplanting/sowing. The over-supply of pre-plant N
potentially results in excessive N loss via either nitrate leaching or ammonia volatilization and
hence diminishes N uptake efficiency
Based on crop N accumulation patterns, it appears that tomato has a relatively

short period of high uptake rates compared to peppers and potatoes (Fig. 6).
Secondly, the homogeneity of the functional relationship for most crops is striking,
despite data being obtained from different studies and production conditions.
Therefore it appears that there are generic uptake trends that hold across crops and
production environments. We thus propose that, especially on soils with limited
nutrient retention and/or with the use of hydroponics, matching fertilizer supply
with crop N requirements may facilitate more efficient N uptake efficiencies. By
applying excessive amounts of pre-plant broadcast applied fertilizer will result in
high residual soil N during the initial 5–6 weeks. Based on the information pre-
sented in Fig. 7, it is evident that crop nutrient requirements during this period are
very low and may only be on the order of 4–12 kg N ha−1. This over-supply of pre-
plant N potentially results in excessive N loss via either nitrate leaching or ammonia
volatilization and hence diminishes N uptake efficiency. Additionally, under water
stress, the over-supply may lower soil water potential and exacerbate the unfavor-
able growth condition.
On the other hand, root growth of especially tomato and pepper during initial
growth is confined to rather small and shallow soil volume (Scholberg 1996). Therefore
overall root N-uptake capacity may limit N uptake during the first 6–8 weeks. The lack
of well-developed root systems with root length densities below 2 cm cm−3 hamper
efficient extraction of soil N thereby reducing N uptake efficiency (Tinker and Nye
2000). However, application of N in excess of crop physiological N requirements still
may be needed to compensate for low extraction efficiencies. This to ensure that N
will not be a limiting factor to crop growth and crop production. With use of tomato
transplants initial tomato root lengths are on the order of 3–4 m plant−1, but this root
length is confined within less than 0.03% of the total soil volume of the production
bed (Scholberg 1996). As soon as the root system expands both vertically and later-
ally, uptake efficiencies increase accordingly (Scholberg et al. 2009). Reduced N
application during crop maturation is also imperative since N requirements during
this growth stage declines significantly. In the case of tomato, N withdrawal and
re-translocation from canopy to fruit during crop maturity may also enhance overall
NUE. It is thus concluded that initial crop N accumulation rates are relatively low.
However, although small rooting volumes and low root densities hamper efficient
nutrient utilization during initial growth, sustained nutrients levels are very much
needed to prevent growth reduction due to N stress. Use of banded slow-release
fertilizers for pre plant application may be a possible solution to address this issue.
6 Nitrogen Uptake Efficiencies
Although tomato, bell pepper and potato have high N demands, their utilization of
fertilizer N typically is rather poor (Table 2). Bell pepper crops generally had the
lowest N recovery (Table 2) as compared to other crops. An more detailed uptake
study showed that for bell pepper N uptake efficiencies increased from 17% at 3
weeks after transplanting to 33% at week 6 where as cumulative seasonal values
were on the order of 45% (Scholberg et al. 2009). Diminished fertilizer recovery
typically occurred with increased application rates, for many crops (Olsen et al.
1993; Joern and Vitosh 1995a, b; Andersen et al. 1999; Tei et al. 1999; Scholberg
et al. 2000a, b; Zotarelli et al. 2009a, b; Zotarelli et al. 2011). Differences in soil
type and cultural practices could explain variability in N recovery in bell pepper and
tomato (Locascio et al. 1985; Andersen et al. 1999). Timing of fertilizer application
also affected uptake efficiency in potato crops (Maidl et al. 2002). However, overall
N recovery and efficiency for these crops seems to be even more impacted by sea-
sonal variation. Although season-to-season fertilizer efficiency was a key source of
variability in FUE, it did not receive strong research emphasis (Locascio et al. 1985;
Roberts et al. 1991; Olsen et al. 1993; Joern and Vitosh 1995a, b; Maidl et al. 2002).
Historically, season was considered a random effect and lower emphasis was placed
to correlate differences in soil temperatures, solar radiation and variation in rainfall
with N uptake efficiency to account for seasonal variation in N-fertilizer uptake
rates (Joern and Vitosh 1995a, b; Thorup-Kristensen and van den Boogaard 1998;
Vavrina et al. 1998).
Nitrogen form and the influence of genotype also affect total uptake of N fertilizer
and fertilizer uptake efficiency. Total uptake of N by the three vegetable crops was
higher under nitrate nutrition when compared with ammonium(Magalhaes and
Wilcox 1983; Marti and Mills 1991; Roberts et al. 1991). Substantial differences in
N-fertilizer recovery and uptake efficiency between cultivars of bell pepper oand
potato occurs. However, genetic selections are commonly based on crop perfor-
mance under optimal fertility conditions. Where as genetic traits also play a
significant role in determining total crop N demand (Kleinkopf et al. 1981; Tei et al.
1999; Alva et al. 2002).
Inefficient fertilizer use by these crops, as evident from low crop N recoveries,
continues to prevail in commercial vegetable production systems. This despite exhaus-
tive management prescriptions for optimal N recommendations, which only effec-
tively address overall crop N needs. Many authors have assessed the effectiveness of
alternative application frequency or methods to increase fertilizer efficiency (Locascio
et al. 1985; Roberts et al. 1991; Olsen et al. 1993; Joern and Vitosh 1995a, b; Vavrina
et al. 1998; Maidl et al. 2002). Use of critical nutrient concentration as determined by
chlorophyll content and/or petiole nitrate concentration(Rhoads et al. 1996) is also
commonly used. However, these approaches still do not provide a proactive or com-
prehensive strategy for improving overall resource use efficiencies including
N-fertilizer. As was pointed out in the previous section, synchronization between fer-
tilizer application and crop N requirement is especially crucial for improving uptake
efficiencies on soils with low water and nutrient retention. Moreover, as environmen-
tal guidelines forces growers to reduce overall N-fertilizer application rates, timing of
fertilizer application will become more critical as a key strategy for enhancing uptake
efficiencies and sustaining crop yields.
Although soil testing for residual N has some merits in other production areas
(Westermann and Kleinkopf 1985), it is usefulness in the southeastern USA for
determining optimal N rate is extremely limited due to N being to readily lost by
leaching in these systems. Especially at high fertilizer rates commonly used for most
vegetable operations, appreciable nitrate leaching may arise. This is especially the
case if poor uptake efficiencies results in high residual soil N concentrations and low
N-fertilizer recovery. Consequently, the leaching of nitrate from vegetable cropping
systems has been a major source of groundwater pollution at a regional level (Tei
et al. 1999; Ramos et al. 2002). Under Florida production conditions crop vigor,
apparent N recovery, and potential denitrification rates differ greatly between drip
irrigated and subsurface irrigated crops. Presence of confining soil layers and/or
perched water tables near the soil surface (“Flatwood” production system) may pre-
vent N contamination of deeper groundwater resources via denitrification(McNeal
et al. 1994; McNeal et al. 1995). In the case of subsurface irrigated tomato, the pre-
dominantly upward flux of water combined with mulched production beds greatly
reduces N leaching and thereby may also enhance apparent N recovery(Scholberg
1996; Zotarelli et al. 2009a, b). However, in these systems excessive rainfalls may
result in rapid rise in perched water table depths. During such events substantial N
losses may occur due to lateral drainage and gravity-induced losses of fertilizer solu-
tions due to their higher specific density (McNeal et al. 1995). In summary, overall N
uptake efficiencies of vegetable crops are relatively low, especially during initial
growth. Improved synchronization between fertilizer application timing and crop N
uptake may thus be needed to sustain productivity if environmental guidelines require
reduction of N inputs for high value crops.
7 Role of Irrigation Management and Root Distribution

on Nutrient Interception
Crop production is intrinsically linked to leaf photosynthesis and canopy size. Water
stress during the vegetative growth can reduce leaf area, plant height, photosynthe-
sis rate and plant development (Kramer and Boyer 1995). Under water-limiting
conditions, crop production increases proportionally to irrigation supply (Baryosef
and Sagiv 1982; Kramer and Boyer 1995). Adequate water supply is thus critical in
maximizing N utilization, crop production and quality of most horticultural crops
(Singh and Arora 1980; Scholberg et al. 2000a, b; Alva 2007). However, on soils
with poor water retention, application of excess water may promote displacement of
nutrients before complete uptake has occurred (Scholberg et al. 2002; Scholberg
et al. 2009). Appropriate irrigation scheduling and matching irrigation amounts
with the effective field capacity of the effective rootzone thus may provide ways to
minimize the incidence of excess N leaching associated with over-irrigation. Using
information from Fig. 6 to match fertilizer supply with crop N demand, fertigation
provides growers with more flexibility. This since it allows them to apply nutrients
more frequently in quantities that closely match short-term crop nutrient requirements.
Injecting N fertigation towards the end of the irrigation cycle may also prevent
immediate N displacement below the soil region with highest root concentration
(Baryosef et al. 1980; Alves et al. 2006; Zotarelli et al. 2008; Zotarelli et al. 2009a,
b; Zotarelli et al. 2011).
The total N uptake capacity of vegetable crops is regulated by their shoot-mass
as was discussed above. However, total crop N accumulation does not govern the
efficiency by which crops take up N. Nitrogen uptake efficiency is typically defined
by N uptake characteristics of the root system and the residence time of N in the
rhizosphere as dictated by rainfall, irrigation, soil type, and presence of mulch
(Scholberg et al. 2002). In studies with spinach (Spinaceaoleracea L.) and kohlrabi
(Brassica oleraceagongyloides L.) it was shown that although both crops had simi-
lar N accumulation rates, kohlrabi required 40% lower soil N concentration when
compared to spinach (Heins and Schenk 1987). In the instance of low uptake
efficiency, relatively high soil N concentrations are necessary to ensure adequate N
uptake and optimal crop tissue N concentrations, which will results in excessive
residual soil N concentrations and thus an increased potential for N leaching. While
some of these processes have been studied for several vegetable crops (Heins and
Schenk 1987; Thorup-Kristensen 1993; Thorup-Kristensen and van den Boogaard
1998; Thorup-Kristensen and van den Boogaard 1999; Cherr et al. 2006), there is a
paucity of data for tomato, potato and bell crops in this regard.
Despite the limited number of existing studies focusing on short-term water and
nutrient uptake dynamics, studies with bell pepper (Scholberg et al. 2009) demon-
strated that improved irrigation scheduling along with a better understanding of
changes in root interception capacity is critical for improving uptake efficiencies..
Rooting depth and width and root length and density of crops are thus key compo-
nents determining potential fertilizer uptake efficiency. Deeper root systems may
more effectively utilize N at greater soil depths and thereby enhance near complete
crop nutrient interception. Studies of leek (Allium porrum L.), white cabbage
(Brassica oleracea L.) and carrot (Daucuscarrota L.) at their optimal rate of fertil-
izer N showed that these crops had different uptake efficiencies. Cabbage had the
highest uptake efficiency and the greatest depletion of residual soil N and both traits
were related to having greater effective root depths (Thorup-Kristensen and Sorensen
1999). In the absence of confining soil layers, tomato was reported to be relatively
deep-rooting across different environment (Jackson and Bloom 1990; Scholberg
1996; Sainju et al. 2000; Zotarelli et al. 2009a, b). However, low recovery of soil N
was reported Jackson and Bloom (1990). Regarding this finding it should be noted
that reported crop yield were very poor (10 t ha−1) compared to those for commer-
cially operated fields. Therefore it appears that the total uptake capacity of the shoot
may not have been achieved. Although there is limited information on the root
uptake efficiency of potato and bell pepper these crops tend to have shallow root
systems. Root excavation studies showed that the rooting depth of potato was gener-
ally no greater than 0.5 m while lateral growth was confined to a soil zone of less
than 0.4 m (De Roo and Waggoner 1961; Lesczynski and Tanner 1976; Munoz-
Arboleda et al. 2006). The soil zone effectively exploited by bell pepper was reported
to be mainly confined to within 0–0.25 m depths in most environments (Hulugalle
and Willatt 1987; Goyal et al. 1988; Zotarelli et al. 2011). The influence of crop
genotypes may also play a key role in root growth as was shown for bell pepper and
tomato (Portas 1973; Leskovar et al. 1989). However, these reports do not capture
the dynamics of root expansion. As a rule of thumb, root elongation is on the order
of 1.25 cm day−1 unless confining soil layers occur (Portas 1973). However, root
growth is also affected by soil temperature, soil moisture, soil bulk density, and crop
N status (Kramer and Boyer 1995). Accurate assessment of root growth and effec-
tive root depth therefore may require development of functional relationships and/
or use of crop growth models(Scholberg 1996).
Depth of rooting and root volume in broccoli (Brassica oleracea L.) was increased
due the presence of high concentrations of residual soil N at greater soil depth (Thorup-
Kristensen 1993). However, work with bell pepper and carrot showed that root volume
diminished with increasing amounts of N being supplied from fertilizer urea or miner-
alized green manure (Goyal et al. 1988; Thorup-Kristensen and Sorensen 1999).
Improved integration of rooting depth in irrigation management strategies also plays an
important role in enhancing uptake efficiency of a crop. Attempts to calculate rooting
depth by using a degree-day approach has been restricted to cauliflower and carrot
crops (Thorup-Kristensen and Sorensen 1999; Thorup-Kristensen and van den
Boogaard 1999). In this regards, base temperatures for potato and bell pepper crops
have been assessed. Potato roots appeared to be unresponsive to changes in soil
temperatures between 9°C and 29°C up to 30 days from emergence. However, at later
development stages they showed exponential growth between 22°C and 29°C (Epstein
1966). In the case of bell pepper, root growth increased exponential between 10°C and
24°C with little change in root growth occurring outside this range (Rylski 1972).
As was stated before, functional relationships are required that will provide
generic relationships of changes in root depth and root length distribution over time.
By integrating this information into experts systems, growers may be able to more
efficiently manage fertilizer resources based on crop development stage, field con-
ditions, environmental guidelines, and production targets as was previously done
for citrus (Morgan et al. 2006). Critical parameters to be include in such information
systems are irrigation and fertilizer practices used, effective root depth as a function
of crop development stage, critical root length, and soil N concentrations required
to ensure maximum N uptake efficiencies and optimal crop performance.
8 Conclusion
This article demonstrated that intrinsically low N-fertilizer recoveries that are
commonplace in high value cropping systems are partly related changes in root
interception capacities and crop N demand over time. It was shown that yield
response to fertilizer application rates for most crops tend to be inconsistent both
within and across locations. Therefore, use of standard recommendations is
undesirable since they pose substantial economic and environmental risks. In this
context, growers may perceive the risks of adverse conditions reducing their
crop yields to be a significant threat. This in combination with the very high tomato
price to fertilizer cost ratio may entice them to use excessively high fertilizer rates
to minimize the risk of yield losses under adverse conditions.
In terms of providing framework for enhanced understanding of processes that
govern N-fertilizer guidelines, we showed that crop N recommendations embody
the interaction between crop physiological N demand and uptake efficiency. In this
manner we aim to make the linkage between genetic production potential, crop N
accumulation, production goals, resource use efficiency, and nutrients management
more transparent. Overall nitrogen uptake efficiency was found to be mainly affected
by inherent genetic traits, environmental conditions and management practices.
Since water and nutrient efficiencies are intrinsically linked, both irrigation and
fertilizer managements need to be optimal to warrant optimal yields and maximum
resource use efficiencies. Increased fertilizer application tends to enhance crop
growth and yield while enhance crop growth will increase overall crop nutrient
demand. However, in many cases excessive N application may delay harvest while
also potentially reduce resource use efficiency, yields and crop quality. Although
total N accumulation differed among crops and locations, temporal N-uptake
patterns were similar for all evaluated crops. Initial crop N accumulation rates and
uptake efficiencies are relatively low. These low efficiencies are caused by small
rooting volumes and low root densities thus hampering efficient nutrient utilization.
However, adequately nutrients soil supply levels are still needed to overcome these
intrinsically low initial extraction efficiencies. Otherwise growth reduction may
cause initial N stress and will result in irreversible crop yield losses. In addition to
this, improved synchronization between fertilizer application timing and current
crop N demand is essential when trying to enhance N-fertilizer efficiencies. This is
especially pertinent for soils with low ater and nutrient holding capacities and
when environmental guidelines dictate reduction of overall N inputs for high value
cropping systems. Developing functional and more generic relationships that
outline changes in crop N demand and effective root depth over time is essential for
the development of expert systems. Use of such systems may allow growers to
more efficiently manage fertilizer resources by more effectively using information
pertaining to crop growth, soil properties, and environmental conditions for devel-
oping farm-specific in-season irrigation and fertilizer management practices.
Acknowledgements This work was partially supported by University of Florida, Institute of

Food and Agricultural Sciences (IFAS); Florida Department of Agriculture and Consumer Services
(FDACS) and Southwest Florida Water Management District (SWFWMD).
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Soil Organic Matter Dynamics and Structure
Nikolaos P. Nikolaidis and G. Bidoglio
Abstract Soil ecosystem functions have significantly deteriorated due to agricultural

intensification with dramatic consequences on carbon loss, loss of soil biodiversity,
erosion, compaction as well as unsustainable use of water and mineral resources.
Sustainable agricultural practices are necessary if we are to face the challenge of food
security while preserving the integrity of soil and aquatic ecosystems. Conservation
agriculture, which is comprised of zero or minimum tillage, carbon amendments
and crop rotations, holds great promise in delivering higher yields, using water and
soil resources in a sustainable manner and increasing soil biodiversity. This article
presents a synthesis of current knowledge on soil ecosystem processes and modeling
with a focus on carbon and nitrogen dynamics and their link to soil structure, and
proposes a conceptual framework for model parameterization capable of predicting
critical soil functions and potential shifts.
We reviewed the dynamics of carbon, nitrogen and soil structure with an empha-
sis in elucidating predominant state variables and the interaction with plants and
food web dynamics. Existing models that simulate the dynamics of organic matter
and structure in soils at various scales were evaluated for their ability to simulate the
functions of soil ecosystem. Current modeling approaches treat carbon, nitrogen
and soil structure for the most part separately without incorporating feedback
mechanisms. The synergistic and antagonistic processes between bacteria and plants
and fungi and plants are partially understood and more importantly the community
lacks the knowledge to predict if and when these processes fail and any related
N.P. Nikolaidis (*)

Department of Environmental Engineering, Technical University of Crete,
73100 Chania, Greece
e-mail: nikolaos.nikolaidis@enveng.tuc.gr
G. Bidoglio
Institute for Environment and Sustainability, Joint Research Centre,
European Commission, Via E. Fermi, 2749, ISPRA(VA) 21027, Italy
176 N.P. Nikolaidis and G. Bidoglio
potential ecosystem shift. A conceptual modeling framework is proposed, developed

along the following three axes: incorporate emerging ecosystem state variables,
account for the ecology of life in soils, and model processes from first principles.
A synthesis of the carbon and nitrogen cycles is suggested in which the dynamics
of the two cycles are interlinked. State variables in soil ecosystem models that
link carbon and nitrogen dynamics with soil structure and the biological community
are recommended. Plant feedback mechanisms with the physical, biochemical
and biotic soil components and the symbiotic relationship between bacteria, fungi,
and plants should be modeled using principles from the ecological succession
theory that would relate the taxonomic structure with function and nutrient fluxes.
A conceptual model of soil structure and soil stability is suggested that links the
soil organic matter sub-model to an aggregation sub-model and a dynamic soil
structure sub-model. The development of new generation soil ecosystem models is
a necessary step to better quantify soil functions, assess possible soil tipping points,
and develop methods to restore soil functions.
Keywords Soil carbon • Soil nitrogen • Soil structure • Model • Feedback

mechanisms
Abbreviations
POM Particulate Organic Matter

SOM Soil Organic Matter
1 Introduction
Food security, our ability to feed the ever increasing human population as well as
satisfy the increases in the standard of living in developing countries, has been one
of the great challenges of our times (UN FAO 2011). The loss of fertile agricultural
land to urbanization, in conjunction to pressures to increase biomass production
for energy use, biofuel, has caused an expansion of agricultural lands to marginal
areas unsuitable for cultivation. Soil ecosystem functions, food and biomass pro-
duction, biodiversity, carbon sequestration, filtering and transformation, raw mate-
rial and landscape, and heritage have significantly deteriorated worldwide during
the past century due to agricultural intensification with dramatic consequences on
carbon loss, loss of soil biodiversity, erosion, compaction as well as unsustainable
use of water and mineral resources (Stagnari et al. 2009; Sapkota 2012). Figure 1
visually illustrates the impacts of tilling and agricultural practices in a olive grove
in Crete, Greece. Similarly, lack of terrace maintenance combined with livestock
grazing have caused failure of the terraces, soil erosion and general deterioration
of the landscape (Fig. 2).
Fig. 1 Impacts of tilling and agricultural practices in a olive grove in Crete, Greece. Top soil has
eroded around the roots of the tree
Fig. 2 Abandoned agricultural terraces in Crete, Greece. Lack of terrace maintenance combined
with livestock grazing have caused failure of the terraces. Soil erosion and degradation leads to
desertification of the land
Conventional agriculture maintain yields by extensive use of fertilizers and

energy (Heller 2009; Tilman 2010) which makes it overall unsustainable. Significant
recycling of NPK nutrients is necessary to sustain global food production (Nikolaidis
2011). Sustainable agricultural practices are necessary if we are to face the challenge
of food security while preserving the integrity of soil and aquatic ecosystems.
Conservation agricultural practices which are comprised of zero or minimum tillage,
carbon amendments and crop rotations (Aune 2012; Stagnari et al. 2009; Sapkota
2012), holds great promise in delivering higher yields, using water and soil resources
in a sustainable manner and increasing soil biodiversity (Saha 2010).
An ecosystem is a dynamic complex of plant, animal and microorganism com-
munities and the non-living environment interacting as a unit (Alcamo et al. 2006).
Ecosystem shifts due to climate and land use changes have abrupt, long-lasting
impacts to ecosystems and should be averted by defining critical thresholds that
provide sufficient response time for management (Biggs et al. 2009). The impor-
tance of improving the forecast reliability of ecosystem state, services and shifts has
been highlighted by Clark et al. (2001) a decade ago and has since been the subject
of many scientific and policy related manuscripts (Schroter et al. 2005; Butler and
Oluoch-Kosura 2006; Nelson et al. 2006; Tallis and Kareiva 2006; Carpenter et al.
2006; Cowling et al. 2008; Steffen 2009; Allen et al. 2010; Kutilek 2011).
There are two fundamental factors that impede our ability to detect and forecast
ecosystem shifts: lack of monitoring data and scientific gaps regarding thresholds in
the function of ecosystems. The limitation in the availability of monitoring data is best
illustrated in the case of land use conversion. Limited chronosequence carbon and
nitrogen data obtained after land use conversion from forest or grassland to crop land
indicate rapid loss of bio-available carbon and nitrogen and deterioration of soil stabil-
ity during the first few years after the conversion (i.e. Olson et al. 2005) suggesting a
significant ecosystem shift. Soil-water-plant models have traditionally simulated con-
ditions where the impacts of land use changes have reached a near-steady state condi-
tion and have not been tested adequately to evaluate the processes during the shifts
(Nelson et al. 2006; Dawson and Smith 2007). Similarly, the models have failed to
predict global shifts such as tree mortality due to climate change or account for the
impacts of inorganic fertilization on soil plant and microbial biodiversity. Our inabil-
ity to predict tree mortality has been associated with gaps in the understanding of the
physiological thresholds and mechanisms of tree mortality as well as the inability to
adequately model plant-soil interactions and feedback mechanisms (Allen et al. 2010).
Similarly, plant species decline has been related to both high (U.K. grasslands –
Stevens et al. 2004) and low-level (Minnesota prairie-grasslands – Clark and Tillman
2008) atmospheric nitrogen inputs suggesting a narrow range of ecosystem thresholds
(Rockstrom et al. 2009) that have neither been fully understood nor the respective
processes incorporated into soil ecosystem models. These examples suggest that a
new generation of soil models should be developed if we are to understand how
to reverse the impacts of land use conversion, redesign agricultural practices, and
quantify the impacts of climate change to terrestrial ecosystems.
During the past decade, there has been an “explosion” in the number of
review articles published in the scientific literature on terrestrial processes and
Soil Organic Matter Dynamics and Structure 179
modeling suggesting a growing demand within the scientific community for a

better understanding of the unifying principles of soil ecology (Fierer et al. 2009).
The objective of this review is to provide a synthesis of current knowledge on soil
ecosystem processes and modeling with a focus on carbon and nitrogen dynamics
and their link to soil structure, and propose a conceptual framework for model
parameterization capable of assessing critical soil functions and potential shifts.
It is not our intent to cite the thousands of articles of primary research, but rather
to focus on review articles that discuss critically and in detail original research
and synthesize the acquired knowledge of these reviews from the view point of
modeling soil ecosystems.
2 Soil Carbon and Nitrogen Dynamics
Carbon and nitrogen dynamics in terrestrial environments have been reviewed

extensively in recent years (i.e. Jastrow et al. 2007; Thevenot et al. 2010; Sinsabaugh
2010; McGuire and Treseder 2010; Kuzyakov 2010; Blagodatskaya and Kuzyakov
2008; Gardenas et al. 2011; Schimel and Bennett 2004; Nannipieri and Eldor 2009;
Geisseler et al. 2010; Knicker 2011; Wu 2011; Szanser et al. 2011). The following
is a synthesis of the literature with an emphasis on elucidating the predominant
state variables of the system and bridging the gap between scientific knowledge and
manner in which organic matter is modeled.
2.1 Carbon Dynamics in Soils
Carbon dynamics in soils are driven by photosynthetically derived plant litter

inputs and organic amendments to the soil system (Fig. 3). Plant litter is physically
fragmented into Particulate Organic Matter (POM) which is composed of lignin,
cellulose and hemi-cellulose (Thevenot et al. 2010). Lignin is connected to cellulose
and hemicellulose in the fiber walls of the vascular system of plants providing
strength and rigidity. Lignin consists of aromatic rings that are chemically resistant
to degradation and thus influence carbon turnover. Cellulose are polysaccharide,
glucose chains, (C6H10O5)n, that comprise the structural material of the cell wall. On
average, one third of all plant matter is comprised of cellulose, although there is
high variability in the composition among vascular plants (i.e. Cotton has 90%
cellulose content and trees 40–50%). Hemicellulose contains several short chain
sugar monomers and binds together with pectin, a structural polysaccharide, to
cellulose to form a network of cross-linked fibres (Thevenot et al. 2010).
Lignin degrades primarily through biotic, aerobic and co-metabolic processes
although a few studies suggest anaerobic biodegradation and abiotic processes
(Thevenot et al. 2010). Saprotrophic fungi and a few bacteria have been shown to
degrade lignin, a process that is catalyzed by unspecific, extracellular enzymes like
Fig. 3 Schematic representation of the combined carbon and nitrogen cycles in soils. The combined
cycles were obtained by taking the point of view of terrestrial biota and follow the fate of organic
matter in soil. The nitrogen cycle emerges as a separate entity after the humification process
lignin or manganese peroxidase. Accumulation and stabilization of lignin in soil has

been observed and attributed to clay-lignin binding, however, this process has not
been elucidated adequately. Rapid turnover of lignin has been observed in agricul-
tural soils suggesting large variability in degradation rates and the existence of two
pools. Thevenot et al. (2010) concluded that there are gaps in our knowledge of the
fate of lignin in soils associated with their stabilization in clay particles and that
lignin should be considered as a state variable in carbon dynamic models.
The major pathways of POM decomposition and creation of humic substances
has been reviewed in depth by Jastrow et al. (2007) and Wardle (2002). POM under-
goes a biochemical alteration into smaller molecules mediated by fungi, bacteria and
enzymes. The products of decomposition are sugars, polyphenols and quinones
which are derived from the microbial decomposition of lignin. The products from
microorganism lysis and released exudates are amino compounds, acids and sugars
as well as nucleic acids and structural cell polymers. Linear chains (>250) of amino
acids form proteins which are essential parts of microorganisms and plants partici-
pating in all processes within their cells. Amino acids are bonded together with
peptide bonds (CO-NH amide moiety) to form the protein chains. Many proteins
are enzymes with specific roles in catalyzing chemical reactions within the cell or
outside the cell which correspond to metabolic reaction and substrate utilization
respectively. Other proteins are involved in signaling processes or as structural
components of the cell. Rillig et al. (2007) classified proteins into two categories,
the detrital proteins that are released after cell lysis and the functional proteins that
include microbial surface-active proteins and extracellular enzymes. Finally, mycor-
rhizal fungi contribute glycoprotein complex molecules to the soil organic carbon
pool through the release of exudates and products of fungal lysis. Condensation
and polymerization reactions between the amino compounds, the sugars and the
quinines and the lignins produce humic substances. Humic substances are associations
of low molecular weight compounds stabilized by hydrophobic interactions and
hydrogen bonds (Sutton and Sposito 2005). Stevenson (1994) proposed four major
humification pathways: (1) the sugar-amine pathway, (2) the polyphenol-quinone
pathway, (3) the lignin-quinone pathway, and (4) the modified lignin pathway. Further
microbial degradation of humic substances results in CO2 which closes the cycle.
The released enzymes mediate key carbon and nitrogen cycle processes involved
in the decomposition and humification of organic matter (Sinsabaugh 2010). Plants
and microorganisms produce enzymes to mitigate oxidative stress, detoxify pheno-
lic compounds and utilize carbon sources. Environmental factors such as oxygen
availability, soil pH, mineral and organic matter composition as well as nitrogen
enrichment regulate the overall enzyme activity in soils. The interactions between
the biological and environmental factors produce positive and negative feedbacks
that control the content of organic matter in soils.
Plant roots release exudates that can “prime” microbial activity (Kuzyakov
2010; Blagodatskaya and Kuzyakov 2008) and the phenomenon is especially
significant in microbial hotspots where the activity is intense. The priming
sequence involves: (1) exudation of organics by the roots, (2) activation of micro-
organism (r-strategists, organisms that have the ability to reproduce quickly) by the
availability of organics, (3) activation of fungi (k-strategists, organisms that have
the ability to compete successfully for resources), (4) production of enzymes by
fungi to catalyze the decomposition of SOM, (5) production of organics and nutri-
ents, and (6) uptake of nutrients by roots (Kuzyakov 2010). Gardenas et al. 2011
identified rhizosphere priming as one of the five most important knowledge gaps in
our understanding of carbon and nitrogen interactions in soils.
Sorption of organic and organo-mineral compounds on clays and oxides is a
significant chemical protection mechanism in the carbon cycle. Organic compounds
bind to mineral surfaces through cation bridges, hydrogen bonding and van der Waals
forces (i.e. Kleber et al. 2007). A number of factors such as pH, redox conditions,
and the characteristics of mineral surfaces and of organic matter affect the degree of
sorption and surface complexation of organics with mineral surfaces. The negatively
charged surfaces of clay minerals bind with the negatively charged organic compounds
through cation bridges (Jastrow et al. 2007). The availability of multivalent cations
plays an important role in the formation of clay-cation-organic complexes. Similarly,
the availability of iron and aluminum oxide surfaces bind organic compounds via
electrostatic forces. Kleber et al. (2007) suggested a conceptual model of organo-mineral

interaction in soils that consists of three layers, the contact zone, the hydrophobic zone
and the kinetic zone. In the contact zone, organic functional groups form inner sphere
complexes by binding directly to the surface. The hydrophobic zone is formed due to
hydrophobic interactions of the organic functional groups in the contact zone and those
in solution and the kinetic zone is formed due to cation bridging and hydrogen bonding
between organics. Chemical protection of carbon is one of the mechanisms that
increases carbon turnover time in soils. The other is physical protection in the soil
aggregates and it will be discussed later in this review.
2.2 Nitrogen Dynamics in Soils
A new conceptual cycle of nitrogen dynamics in terrestrial systems is emerging based

on the research conducted during the past two decades (Fig. 1). The following is a
synthesis of several recent review papers (Schimel and Bennett 2004; Nannipieri and
Eldor 2009; Gardenas et al. 2011; Geisseler et al. 2010). Soil organic nitrogen being
part of the humic substances is comprised of proteins, peptides and amino acids (40%),
amino sugars (6%), heterocyclic N compounds (ie. Pyrines, 35%), and ammonia (19%)
(Schulten and Schnitzer 1998). The traditional view of the nitrogen cycle is that soil
organic nitrogen is being mineralized to ammonia and ammonium ion due to micro-
bial activity in the soils (Mineralization-Immobilization-Turnover route). Ammonium
is then converted to nitrite and then to nitrate through the microbial assisted nitrification
process. Denitrification converts nitrate to nitrogen gas and nitrous oxide. Microbial
cell synthesis requires primarily ammonium uptake creating an immobilization mecha-
nism, while plants uptake nitrate and ammonium for the development of plant tissue.
Schimel and Bennett (2004) reviewed the literature of mineralization studies and sug-
gested the need to revise the conceptualization of the N cycle by including the enzyme
catalyzed de-polymerization of soil organic nitrogen into light molecular weight Org-N
such as amino-acids and amino-sugars. They also suggested that plants uptake directly
these monomeric Org-N compounds corresponding to an alternative direct route
(Schimel and Bennett 2004; Wu 2011). This pathway of N immobilization appears to
be very important in low-N systems. This uptake is facilitated by mycorrhizae (Manzoni
and Porporato 2009; Wu 2011). Gardenas et al. (2011) have identified plant utilization
of organic N priming as one of the five most important knowledge gaps in our under-
standing of carbon and nitrogen interactions in soils. Many field studies suggest that
organic N utilization is a ubiquitous capacity of plants, being more important in cool
and wet environments than hot and dry (Gardenas et al. 2011; Jamtgard et al. 2010;
Ramirez et al. 2010). Nannipieri and Eldor (2009) focused on reviewing the mineral-
ization-immobilization processes of N in soil and attempted to relate N turnover to the
biota that conduct the transformation. They extended the view of Schimel and
Bennett (2004) by identifying three pathways of org-N transformation: (1) direct
mineralization of org-N to ammonium by microorganisms, (2) release of org-N due to
micro-organism lysis and exudates, and (3) excretion of ammonium by bacterial-grazing
protozoa and nematodes. Two additional processes have been suggested as being
important in the microbial N transformation in soils: Nitrifier denitrification which is

the oxidation of ammonium to nitrite followed by reduction of nitrite to nitrogen gas
and anaerobic ammonia oxidation or Anammox (oxidation of ammonium to nitrite fol-
lowed by reduction of nitrite to hydroxylamine; hydroxylamine then reacts with ammo-
nium to form hydrazine which is oxidized to nitrogen gas and the released electrons are
used to reduce nitrite). Geisseler et al. (2010) reviewed the pathways of nitrogen uptake
by microorganisms in terrestrial environments and examined the relative importance
between the direct route of org-N uptake versus the mineralization-immobilization-
turnover route. Three factors were identified that determine the relative importance of
the two routes in various ecosystems: the form of available N, the source of C, and the
availability of N relative to C.
A synthesis of these reviews resulted in the following terrestrial nitrogen cycle. POM
is decomposed by saprotrophic fungi to soil organic nitrogen. Microorganisms uptake
and decompose POM. The products of decomposition contribute to the soil organic
nitrogen pool as well as products from microorganism lysis and released exudates. Plant
roots on the other hand release exudates that can “prime” microbial activity. Mycorrhizal
fungi contribute to the soil organic nitrogen pool through the release of exudates and
products of fungal lysis. Microorganism and fungal exudates include enzymes that
catalyze the de-polymerization of soil organic nitrogen to amino-acids and amino-sugar
type compounds. These compounds can be up-taken directly by the microorganisms and
the mycorrhizal fungi/plants or follow the mineralization-immobilization-turnover
process to ammonium. The nitrification process converts ammonium to nitrite and
nitrate while denitrification nitrate to nitrogen gas. Ammonium and nitrate can then
be up-taken by both microorganisms and mycorrhizal fungi/plants. To complete the
list of nitrogen transformations in soils, one could add the two processes of ammonium
conversion to nitrogen gas, nitrifier denitrification and anaerobic ammonia oxidation.
The interaction of the various nitrogen compounds with mineral surfaces should also
be taken under consideration because it plays a role in the regulation of nitrogen trans-
formation in soils by limiting its availability. Plant and microorganism exudates include
protein N and peptide N which can absorb readily to clays due to their hydrophobic
domains. Jamtgard et al. (2010) found that the concentration of bound amino acids in
unfertilized soils were 50 times higher than the free amino acids in solution as well as
higher than the ammonium and nitrate concentrations suggesting the importance of soils
in regulating the availability of enzymes for soil organic matter decomposition.
2.3 Impact of Bacteria, Fungi, Invertebrates and Plants

on the C and N Dynamics
The role of bacteria and fungi in the dynamics of carbon and nitrogen cycles has
already been discussed. They utilize and decompose organic matter and produce
enzymes that assist in the humification process. In addition, they metabolize carbon
and nitrogen for biomass formation. The lysis products of bacteria and fungi are
very different in nature. Fungal cells are composed of complexes such as melanin
and chitin (amino sugars) whereas bacterial cells primarily of phospholipids and
peptidoglycan (Lorenz et al. 2007). Melanin and chitin do not degrade easily, while
phospholipids are metabolized rapidly by other bacteria and grazers. Soil invertebrates
such as protozoa and nematodes contribute significantly to plant litter decomposition
and microbial population control through grazing (Jastrow et al. 2007). A detailed
discussion of soil food web structure and dynamics is given by Wardle (2002).
A conceptual representation of the symbiotic relationship between ectomycorrhizal
fungi and plants in a forested ecosystem has been reviewed by Courty et al. (2010).
In addition to participating in the carbon and nitrogen cycles, mycorrhizal fungi
contribute to soil functions by linking trees together through common mycorrhizal
networks. Mycorrhizal community taxonomical structure and growth depends on the
quantity and quality of tree photosynthetic activity, soil water, nutrient content and
mineral composition. The taxonomic structure and the soil properties determine the
functional structure of the mycorrhizal community which in turn affects the mineral
nutrition of the tree. The nutritional status of the tree controls its photosynthetic
activity, closing in this way the feedback mechanism. Recent research (Bonfante and
Genre 2010) has identified selective transporters located in the fungal membrane
that facilitate the translocation of carbon, nitrogen and phosphorous from the soil to
the plant tissue, supplying the plant with nutrients. Fungi produce hyphal branching
in the direction of root exudates in search of new root colonization suggesting direct
signaling communication. Intracellular and intercellular hyphae colonization of
the root provides large surface area for nutrient exchange. Plant growth-promoting
bacteria that colonize the rhizosphere and the plants (endophytic colonization)
compliment the role of fungi in the plant nutrition cycle. In addition, the endophytic
bacteria have been shown to be capable of degrading contaminants (phytoremediation
of organic pollutants) and contributing in this way to the defense mechanisms of
plants (Compant et al. 2010; Cheng et al. 2010). Bacteria use extracellular (quorum-
sensing) and intracellular signaling to assess the external conditions and react to
environmental changes (Camilli and Bassler 2006). Similar biochemical signaling
exists between plant and microbe interactions (Baker et al. 1997). Feedback mechanisms
in the plant-soil system (in terms of the physical, biochemical and biotic components
of the soil) were reviewed by Ehrenfeld et al. (2005) in an attempt to critically appraise
the system and identify new research directions (Table 1). There is strong evidence
of feedback mechanisms in the plant-soil system which become more apparent in
extreme environments and systems involving trophic interactions. There is a need to
further study the feedback between soil structure and plant growth as well as the
role of plant-microbial plasticity in enhancing or attenuating feedbacks.
2.4 Modeling of Carbon and Nitrogen Dynamics
In general, carbon models divide SOM into several pools one of which is the soil
microorganism pool. Carbon in these pools is degraded as a first order process. This
approach is also used in eco-hydrological models (i.e. SWAT, Debele et al. 2008 or
SWIM, Post et al. 2007) that simulate carbon, nitrogen and phosphorous dynamics at
the watershed scale. They have been using a simplified parameterization of carbon
Table 1 Summary of feedback mechanism in the plant-soil system (Ehrenfeld et al. 2005)
Plant feedbacks with physical soil components
Soil moisture feedbacks
Capacity of plants to alter the distribution of water in soil
Plant adaptation to water stress
Arbuscular mycorrhizae promote hydraulic lift, modify soil hydrophobicity, change water
routing
Soil aggregates
Roots contribute to SOM input
Root exudates stimulate soil aggregation
Temperature
Shading by litter and aboveground biomass
Changes in albedo
Plant feedbacks with biochemical soil components
Soil acidity
Plant adaptation to pH involve complex biochemical, physiological and mutualistic pathways
Small pH changes affect plant growth
Plants participate in the acidification of soil
Cations
Metal hyper accumulation feedback
Plants affect soil cation concentration through mineral weathering, uptake and redistribution
Plants affect soil redox conditions and redox sensitive metals
Carbon and nitrogen
Litter input affects microbial mineralization
Feedbacks that regulate the supply of nutrients
Plant affects carbon and nitrogen storage in soil
Plant feedbacks with biotic soil components
Microbial community structure feedback
Sensitive to plant species
Rhizosphere microbes respond to plant exudates due to plasmids with genes
Quorum sensing mechanisms for regulating expressions of trait
Plant protein can disrupt the plant-microorganism “dialogue”
Mutualistic feedback
Communication through molecular and genetic feedbacks
Root exudates promote the growth of disease suppressing bacteria
Mycorrhizal mutualism operates at the level of individual plants and affects plant growth
Feedbacks with pathogens, parasites and hervivores
Attack plant tissue
Affect tree seedling
Promote the invasion of exotic species
Feedbacks with Invertebrates
Part of feedback loops linking carbon, nitrogen, microorganisms and plants
Soil food web affects plant growth
Invertebrate predation on soil microbial biomass affects nutrient mineralization rates
and nitrogen in the soil that consists of recalcitrant and labile organic pools as well as
the mineral pool. The use of the first order exponential model to describe the dynamics
of carbon degradation even though it “fits the data” in many cases, it does not accurately
reflect the dynamics of the process and alternate models have been proposed. These
alternate models are based on the assumption of time-variable decomposition rate
(Rovira and Rovira 2010) or the incorporation of logistic functions, recognizing the
existence of competitive strategies in soil (Gillis and Price 2011). Carbon and nitrogen
mineralization is considered the dominant process that drives the system. The most
comprehensive parameterization regarding soil formation is presented in the SoilGen1
model (Finke and Hutson 2008). SoilGen1 is an integration of a transport and spe-
ciation of inorganic anions model and a carbon cycle model. The one dimensional
transport Richards’ equation for transient flow is coupled with heat flow and the
advective-dispersive equation for solutes. Major anions and cations are simulated and
chemical speciation determines the precipitated and exchanged phase based on ther-
modynamic considerations. The carbon cycle model considers five carbon pools, the
resistant and decomposable plant material, the microbial biomass, the humified organic
matter and the inert organic matter. Mass changes due to bioturbation, organic carbon
and precipitates are used to calculate changes in bulk density and porosity and thus soil
evolution. In the models described above, the microorganism communities have been
modeled in terms of biological carbon content using a static physiological state.
Even though modeling of several bacteria and fungi communities shifting in com-
position or transition from an active to dormant state is still at its infancy, the approach
holds great promise and should be further developed (Ingwersen et al. 2008).
To more accurately account for the turnover of carbon in arable soils, Kuka et al.
(2007) developed Carbon turnover In Pore Space (CIPS) model. The model assumes
that the biological activity is not distributed evenly in the pores which results in
uneven oxygen distribution and differences in decomposition rates. It considers
three pore sizes, the micro-pores, meso-pores and macro-pores and four carbon
pools corresponding to fresh, active, refractory and dissolved organic matter. In this
way, soil carbon turnover time was related to physical measures of the soil system.
Modeling of carbon and nitrogen dynamics in terrestrial environments has also
been reviewed extensively in recent years (i.e. Falloon and Smith 2000; Manzoni and
Porporato 2009; Batlle-Aguilar et al. 2011; Shibu et al. 2006; Minasny et al. 2008).
Fallon and Smith (2000) reviewed 98 agro-ecosystem models in order to identify the
shortcomings in modeling refractory soil organic matter and set priorities in research
areas for future development. They concluded that processes of refractory SOM for-
mation and protection are not clearly understood and models were limited in their
ability to predict refractory SOM content and turnover because refractory SOM pools
cannot be directly related to field measurements. Models need to account for carbon
turnover with depth as well as for carbon stabilized on charged surfaces of iron oxides
and clays. Manzoni and Porporato (2009) reviewed and classified 250 biogeochemi-
cal models in terms of their mathematical approaches to soil carbon and nitrogen
dynamics. In addition, they analyzed the relationship between model structure and
the temporal and spatial scale of its respective application. They identified the
following theoretical gaps: (1) mechanistic and scale dependent description of micro-
bial mass and activity using dedicated state variables, (2) link decomposer activity
and metabolism, nutrient availability, vegetation growth and climate dynamics

using stoichiometric links, (3) soil food web dynamics instead of aggregated variables,
(4) mechanistic and rigorous representations of small-scale processes that account
for spatial heterogeneity, (5) physical processes that affect soil structure (tillage,
wetting and drying cycles and aggregate turnover) in the biogeochemical models,
and (6) need to develop novel modeling approaches for scaling up pore scale dynam-
ics to observational scales. Batlle-Aguilar et al. (2011) reviewed carbon and nitrogen
models for their ability to simulate land-use change conditions. The most common
features of the mechanistic compartment based carbon and nitrogen models were the
number of C and N pools, plant growth parameterization, parameterization of physi-
cal soil parameters, simulation time step, interconnectivity of C and N cycles and
simulation of hydrology and soil temperature. An example was used to illustrate the
predictive ability of a model to simulate scenarios of carbon and nitrogen turnover in
forested and agricultural land uses in different climatic conditions. They concluded
that such models are a promising tool in the design of new soil practices and predicting
the effects of the practices on soil fertility. Shibu et al. (2006) reviewed 20 process-
based SOM dynamics simulation models in terms of their ability to simulate long
term carbon and nitrogen dynamics in rice-based cropping systems. They concluded
that no existing SOM model simulates properly lowland anaerobic conditions or
alternate flooded and non-flooded conditions. It is suggested that the biochemical
characteristics of SOM should be modeled explicitly in relation to their effect on
yield. Finally, Minasny et al. (2008) reviewed the applicability of existing quantitative
models for pedogenesis and suggested that an integration of state factor empirical
models and energy and mass balance models could provide the analytical framework
to model how soil horizons and profiles evolve in the landscape.
3 Soil Structure Modeling
Soil structure and in particular soil particle aggregation formation stability and turn-
over play a catalyzing role in soil erosion, nutrient cycling and soil fertility. There
are tens of literature review manuscripts published during the past decade that
review scientific work that supports overwhelmingly the relationship between soil
structure and soil fertility (i.e. Falloon and Smith 2000; Manzoni and Porporato
2009; Batlle-Aguilar et al. 2011; Abiven et al. 2009; Jastrow et al. 2007; Bronick
and Lal 2005; Ehrenfeld et al. 2005). The following is a synthesis of the literature
with an emphasis on elucidating the predominant factors affecting soil structure, its
relation to the carbon and nitrogen cycles and modeling approaches.
3.1 Factors Affecting Soil Structure
Many factors play a significant role in the formation of soil aggregates and their
stability such as the primary particles (clays-silt-sand) and their surface charges,
organic matter available for decomposition (hydrophobic nature – attached to
surfaces); bacterial biomass that utilizes the organic matter and produces exudates
that catalyze the breakdown and transformation of organic matter; plant roots that
also produce exudates, organic matter for decomposition and control the availability
of water in the soil system, and fungi and other organic fauna such as invertebrates
and worms (Ehrenfeld et al. 2005). There are two basic mechanisms of aggregate
formation (Abiven et al. 2007; Abiven et al. 2009). The first mechanism proposes
that microaggregates (<250 mm) consist of organic molecules that are bound to
clays and polyvalent cations to form an organo-mineral complex. Organo-mineral
complexes then combine to form macroaggregates. The second mechanism pro-
poses that macroaggregates are formed around particulate organic matter (POM).
As POM is decomposed bacterial colonies form and bind clay particles with their
exudates. Particle enmeshment to the macroaggregate is facilitated by roots and
fungal hyphae that release exudates that bind particles. In this way, stable microag-
gregates are formed within the macroaggregate and as decomposition (carbon turn-
over) progresses they are released. The stability of aggregates and their turnover
greatly depends on the quantity and quality of organic matter and plant litter added
to the soil (Abiven et al. 2007; Abiven et al. 2009). However, abiotic factors such as
soil texture are more important for determining soil aggregation than biotic factors
(Barto et al. 2010). Earthworms contribute greatly to soil aggregation by utilizing
plant litter together with soil, passing it through their gut and excreting casts (Six
et al. 2004). Casts mediate both microaggregate and macroaggregate formation in
soils the stability of which depends on the quality of organic material consumed by
earthworms. In addition, earthworms contribute to aggregate formation due to bur-
rowing activities where they deposit mucus on the burrow walls which together with
the soil clays can form a stable structure (Six et al. 2004). Roots influence soil
aggregation through entanglement of soil particles with the mycorrhizal-root sys-
tem as well as through root exudates (Jastrow et al. 2007; Six et al. 2004). Table 2
presents an overview of the factors (Bronick and Lal 2005) that must be considered
in the formulation of soil stability and soil fertility models. Microaggregates and
macroaggregates provide a physical protection from decomposer activity to the soil
organic carbon and nitrogen incorporated in them and influence carbon and nitrogen
turnover in soils. Decomposer activity is limited in the aggregate structures due to
oxygen and water availability, and enzyme accessibility (Jastrow et al. 2007).
Figure 4 is a photograph of macroaggregates developed in an uncultivated olive
grove in Crete, Greece. Fungal hyphae and roots participate in the aggregate forma-
tion and they are visible in the upper part of the photograph.
A critical review of the scientific literature (Bronick and Lal 2005; Allen et al.
2010; Six et al. 2004; Alaoui et al. 2011) suggests that even though there has been
significant progress in the understanding of soil processes, a comprehensive under-
standing of how these factors contribute to soil aggregation is lacking. For instance,
a fundamental framework that would explain why some crops are more effective in
structural development than others or how insecticides and fungicides affect aggre-
gate formation is lacking. The link between soil structure and agricultural practices
should be approached from a fundamental perspective if we are to make progress in
developing sustainable crop management systems.
Table 2 Factors affecting soil structure (Bronick and Lal 2005)

Climate and exogenous factors
Climate and Terrain
Temperature and moisture regimes (wet-dry and freeze-thaw cycles) affect biological
activity
Wet-dry cycles affect particle aggregation and stability
Wet-dry cycles affect carbon protection
Elevation, aspect and slope affect vegetation and erosion
Soil properties
Soil texture
Plant adaptation to pH involve complex biochemical, physiological and mutualistic
pathways
Small pH changes affect plant growth
Plants participate in the acidification of soil
Cation exchange capacity
Metal hyper accumulation feedback
Plants affect soil cation concentration through mineral weathering, uptake and redistribution
Plants affect soil redox conditions and redox sensitive metals
Soil pH
Litter input affects microbial mineralization
Feedbacks that regulate the supply of nutrients
Plant affects carbon and nitrogen storage in soil
Agents of aggregation
Organic and inorganic carbon
Creates hot spots of aggregation
Composition affects aggregate stability and turnover
Increases microbial biomass
Particulate organic carbon acts as the nuclei for aggregation
At low organic carbon levels, carbonates decrease carbon mineralization
High levels of carbonates in silt decreases aggregation and stability
Clays
The degree of aggregation, stability and carbon turnover depends on the content of clay in
the soil
Clay mineralogy and surface properties affect soil development
Affects the rate of decomposition of organic matter
Cations
Improve soil structure through cationic bridging with clays and organic matter
Biotic influences
Plants and plant roots
Water stable aggregation is related to the composition of plant residue
Root exudates act as binding agents
Roots enmesh particles and improve stability
Microorganisms (bacteria and fungi)
Fungal and bacterial exudates (polysaccharides) act as binding agents
Fungal hyphae enmesh particles and improve stability
Soil fauna (insects, anthropods, earthworms, nematodes and termites)
Attack plant tissue
Affect tree seedling
Fig. 4 A photograph of
macroaggregates from an
uncultivated olive grove
in Crete, Greece. Fungal
hyphae and roots are
visible in the upper part
of the photograph
3.2 Soil Structure Modeling
During the past decade, there were five papers that dealt explicitly with soil aggregate
modeling (De Gryze et al. 2005; Plante et al. 2002; De Gryze et al. 2006; Abiven et al.
2008; Malamoud et al. 2009). The models can be classified into four classes: empirical,
mechanistic, deterministic soil aggregate models and deterministic soil structure models.
3.2.1 Empirical Models
De Gryze et al. (2005) developed two empirical models, a linear regression and a
3-parameter sigmoidal curve model to fit the rate of increase of water stable aggre-
gates due to POM addition. Abiven et al. (2008) following a different approach devel-
oped an empirical model where Monnier’s conceptual description of variations in
aggregate stability for different organic substrates was formalized and applied to field
conditions. The following log normal function was fitted to the field data: AS(t) = A
exp{−0,5 exp[ln(t/C/B)]2} where A is a function of the lignin content, B of the
polysaccharide content and C of the cellulose and hemicelluloses content. In order
to apply the model to field conditions, factors that account for the effect of soil
moisture, soil temperature and soil N availability on carbon decomposition were
used to adjust the aggregate stability function. These models represent the first attempt
to model aggregate stability dynamics after the incorporation of organic residues.
3.2.2 Mechanistic Models
Plante et al. (2002) assumed four water stable aggregate classes (<1 mm, 1–2 mm,
2–4 mm, and >4 mm) and described the flows between the classes using first order
kinetics. They assumed six potential aggregation (outward flows) and six break-
down (inward flows) movements between the four classes as part of the “fully
developed” conceptual model. The model was calibrated using data for two soils
after tilling. De Gryze et al. (2006) used this model to calculate micro-aggregate and
macro-aggregate turnover times. The soil sample was homogenized with rare earth
powders as tracers to test the assumptions involved in modeling. Mechanistic mod-
els were shown to provide valuable information on the microaggregate and macro-
aggregate stability and turnover, confirming the assumptions that microaggregate
turnover is higher than the macroaggregate one.
3.2.3 Deterministic Soil Aggregate Models
De Gryze et al. (2005) wrote a mass balance on the rate of soil aggregation by
assuming that the rate of aggregation is proportional to mass of non-aggregated soil
minus the mass of aggregate breakdown that is proportional to the aggregate mass:
dM/dt = fU-bM where M is the macroaggregate content of the soil, U is the non-
aggregated soil content, f is the rate of formation and b is the rate of breakdown.
Mass conservation is assured by M + U = 100%. The solution of the mass balance is
an “exponential increasing to maximum” curve. As an alternative to this model, the
authors assumed that the rate of aggregate formation is proportional to microbial
activity in the soil, C. f = kC(t−t0), where t0 is a time lag between microbial activity
and aggregate formation. A measure of microbial activity can be considered the
respiration rate.
3.2.4 Deterministic Soil Structure Models
Malamoud et al. (2009) modified the RothC-26.3 model (Coleman and Jenkinson
1999) to accommodate the concept of size hierarchy for the physical protection of
aggregates and developed aggregation and porosity sub-models in order to model
aggregate fractionation and carbon content, aggregate turnover time and changes in
the porosity of the soil. The new model, named Struc-C, was then used to model data
from an agricultural land and a woodland located in temperate climate with significant
oceanic influence (UK) in the first and to a lesser extent (Australia) in the second (Paul
et al. 2002). The model defined three aggregate fractions with corresponding three
aggregate carbon types and complexed clay distributed in these three fractions assum-
ing constant partitioning with soil organic carbon for each fraction. Once the organo-
mineral aggregates have been calculated, the porosity submodel estimates the bulk
density of each of the three aggregate types assuming pyramidal, tetragonal and cubic
packing respectively and then the new porosity of the soil. The major limitations of
the model as outlined by its authors relate to the following: (1) the relation between
input residue and soil organic carbon is not linear and the quality of the input is not
addressed, (2) geochemical factors that influence soil structure such as type of clay,
iron oxide sorption, pH etc. are not included in the formulation, and (3) aggregate
parameters do not vary with space and time. Even though these are significant limita-
tions of the model, Struc-C is the most comprehensive model regarding soil aggregate
stability and turnover as well as soil structure in the scientific literature thus far.
So far models have aimed at describing the rates of aggregate formation and
turnover. A number of processes that are significant and affect aggregate formation
and stability have not been included (Malamoud et al. 2009; Ehrenfeld et al. 2005)
such as:
• Explicit formulations of aggregate stability as a function of residue type
• Cations and charged surfaces explicitly (acid–base chemistry, CEC, pH)
• Soil moisture impacts and wet-dry cycles
• The concept of Water Stable Aggregate
• Plant and bacterial exudates
• POM composition and decomposition products (lignin, cellulose, polysaccharides)
• Plant and fungi impact to Water Stable Aggregates due to moisture re-distribution
• Antagonistic and synergistic effects on Water Stable Aggregates
• Feedback mechanism
Bacterial communities, soil fauna and plants participate in both synergistic and
antagonistic ways in “building” (aggregate formation) their soil micro-environment
as well as in the “quality” (aggregate stability) of the construction. If one considers
that the rate of microbial reproduction is at the 20 min time-scale, then the life-cycle
of the aggregates which corresponds from weeks to months is very long. In order
to model aggregate stability, the soil ecosystem state variables should include
the “building materials” such as lignin, cellulose, polysaccharides, acid–base and
interfacial chemistry concepts and the synergistic and antagonistic effects of the
terrestrial ecology fauna.
4 Future Directions of Soil Ecosystem Modeling
The complexity of the soil-water-plant-organism system is overwhelming, and even

though ecological and biogeochemical research has made significant strides during
the past decades, we still have significant knowledge gaps in understanding the
fundamental functioning of the system (Gardenas et al. 2011). This has been stressed
in a special issue of the journal of Science, where soils have been identified as “the
final frontier” (Science 2004). There is an emerging consensus that the diversity of
life in soil is a key factor determining soil functions and the life-sustaining pro-
cesses (Crawford et al. 2005). The synergistic and antagonistic processes between
bacteria and plants and fungi and plants are partially understood and more impor-
tantly the community lacks the knowledge to predict if and when these processes
fail and any related potential ecosystem shift. Current modeling approaches treat
carbon, nitrogen and soil structure for the most part separately without incorporating
feedback mechanisms. For instance, recent research (Ramirez et al. 2010) suggests
that inorganic nitrogen fertilization directly affects microbial respiration rates by up
to 60% regardless of the form of nitrogen addition and that fertilization influences
the C and N pools in the aggregates (Chen et al. 2010). Such common perturbation
of soil systems as agricultural fertilization, cannot be modeled properly because
the cause and effect relationships have not been determined. A revised modeling
framework is necessary to be developed that would synthesize current understanding
of soil functions. We propose this modeling framework be developed along the
following three axes:
Emerging ecosystem state variables – A synthesis of the carbon and nitrogen
cycles emerging from this review is presented in Fig. 1. Carbon and nitrogen dynamics
are interlinked and can only be separated after the production of humic substances
by condensation and polymerization reactions (Batlle-Aguilar et al. 2011). The
enzyme catalyzed de-polymerization of humic substances separate the nitrogen
from the carbon cycle by creating light molecular weight organic nitrogen com-
pounds that can be mineralized into ammonium and follow the remaining reactions
in the known nitrogen cycle. State variables in soil ecosystem models should be
selected in a way that link carbon and nitrogen dynamics with soil structure and the
biological community. We propose that carbon and nitrogen state variables be linked
directly with the composition and quality of plant litter and organic matter input to
soil. Soil ecosystem state variables should be comprised of the following:
• The components of the decomposition of particulate organic matter such as lignin,
quinone, polyphenols, sugars, amino compounds, glycoprotein, humics, light
molecular weight organic N compounds, ammonium ion, nitrite and nitrate,
• The biomass of the soil food web such as microorganisms, saprotrophic fungi,
mycorrhizal fungi and other terrestrial species and
• The enzymes produced by microbial and fungal lysis and root exudates.
The adaptation of the proposed state variables in the new generation soil models
would be a strategic decision by the soil science and agricultural engineering com-
munity because it will require extensive modifications in the design and execution of
field studies since current approaches do not measure specifically for these variables.
Modeling ecology of life in soils – Biological diversity dynamics and their interaction
with the physical and chemical microenvironments are emerging as the principal fac-
tor determining soil functions (Barot et al. 2007; Wu et al. 2011; Zhang and Xu 2008).
Plant feedback mechanisms with the physical, chemical and biotic soil components
are critical processes especially in extreme environments that should be considered in
soil ecosystem models. The symbiotic relationship between bacteria, fungi, and plants
should be modeled using principles from the ecological succession theory that would
relate the taxonomic structure of the various species with their respective function in
the ecosystem and the necessary nutrient fluxes to maintain it. It is necessary to view
the ecology of life in soils as an “intelligent” living system within which each species
actively functions towards building its own niche, understanding the symbiotic and
Fig. 5 Schematic representation of a soil structure conceptual model
antagonistic relationships with other species in terms of their own survival and the
survival of the system and participate fully in sustaining soil ecosystem functions
(Gillis and Price 2011). It is also important to assess the capability of soil microbial
communities to adapt to changing environmental conditions and the dynamics of
turnover at different time scales (Schmidt et al. 2007). Soil ecologists and biologists
should join their experiences and propose modeling approaches that would capture
the complexity of the life in soils.
Modeling from first principles – Modeling soil structure and relating soil stability
to carbon and nitrogen dynamics would require a first principles approach where the
interaction within and between the various components of the system (clay parti-
cles, organic matter, terrestrial organisms etc.) are modeled using the fundamental
principles that govern the system. Figure 5 presents a conceptual model of soil
structure and soil stability similar to the one developed by Malamoud et al. (2009).
The SOM sub-model (Fig. 1) should be linked to an aggregation sub-model and a
dynamic soil structure sub-model. The particle distribution of the original mineral
matrix will undergo physical aggregation due to charged surfaces that can be
described and modeled with well developed geochemical principles. On the other
hand, SOM fractions that will parameterize the SOM sub-model will partition to
clay particles and clay aggregates due to their hydrophobic nature. Microbial
decomposition will initiate the aggregate formation and breakdown mechanism as
described earlier. Plant roots and soil biota will contribute in the formation of the
aggregates and the quality of processed plant litter will determine the rate of aggre-
gation and the strength of the aggregates. The aggregate fractionation should then
be evaluated in terms of its stability using empirical meta-models derived from
water stable aggregate studies and the physical changes to soil structure including
changes in bulk density, porosity and change of volume.
5 Conclusion
In this review, a conceptual modeling framework is proposed, developed along the

three axes: incorporate emerging ecosystem state variables, account for the ecology
of life in soils, and model processes from first principles. In particular, the following
are proposed:
• The dynamics of the carbon and nitrogen cycles to be interlinked,
• Soil ecosystem models should be developed with state variables that link carbon
and nitrogen dynamics with soil structure and the biological community,
• Principles from the ecological succession theory that relate taxonomic structure
with function and nutrient fluxes should be used to model plant feedback mecha-
nisms with the physical, biochemical and biotic soil components and the symbi-
otic relationship between bacteria, fungi and plants, and
• A conceptual of soil structure and soil stability model is suggested that links
dynamically soil organic matter to soil aggregation and structure.
The development of a new generation of soil ecosystem models is a necessary
first step in order to quantify soil dynamics, understand soil tipping points, identify
soils at threat and develop methods to restore soil functions.
Acknowledgements Funding for this work was provided by the EU FP7-ENV-2009 Project
SoilTrEC “Soil Transformations in European Catchments” (Grant #244118). This work was con-
ducted at the Institute for Environment and Sustainability of the Joint Research Centre (JRC) of the
European Commission. Professor Nikolaidis is grateful for the Technical University of Crete
financial support of his sabbatical leave at the JRC.
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Plant and Animal Breeding as Starting Points
for Sustainable Agriculture
Gerhard Flachowsky, Ulrich Meyer, and Manfred Gruen
Abstract The present century is characterized by a growing world population and

a higher need for food and natural resources such as water, arable land, fuel and
minerals as well as by growing emissions of greenhouse gas. Agriculture contribute
to a certain degree to resource needs and pollution. Therefore this article shows the
potentials of plant and animal breeding for the most efficient use of non-renewable
resources and consequences for emissions. The efficient production and use of
phytogenic biomass is a challenge for sustainable agriculture. Plant breeding in the
traditional way and green biotechnology are the starting point for the whole food
chain and many other processes based on phytogenic biomass. High and stable plant
yields with low external inputs of limited resources such as water, fuel, land and
minerals are major objectives of plant breeding. Plants must be more resistant
against biotic and abiotic stressors. Plants should use very efficiently unlimited
natural resources such as sun energy, nitrogen and carbon dioxide from the air.
For animal nutrition a lower concentration of toxic substances in plants is more
important than higher yields of components of nutritive value. Enough high quality
phytogenic biomass for animal feeding and animal breeding can also contribute to
a more efficient conversion of feed into food of animal origin and lower emissions.
One of the main objectives of animal breeding is a high dry matter intake in order to
have more energy for animal yields and to reduce the portion for maintenance of
animals. In general plant and animal breeding may be considered as the starting
G. Flachowsky (*) • U. Meyer

Institute of Animal Nutrition, Friedrich-Loeffler-Institute (FLI),
Federal Research Institute for Animal Health, Bundesallee 50,
Braunschweig 38116, Germany
e-mail: gerhard.flachowsky@t-online.de
M. Gruen
Food GmbH – Analytic Consulting, Orlaweg 2, Jena 07743, Germany
e-mail: m.gruen@food-jena.de
202 G. Flachowsky et al.
points for sustainable agriculture in the future. Examples for a more sustainable
agriculture are shown in this article.
Keywords Limited resources • Human nutrition • Carbon footprints • Plant breeding

• Animal breeding • Animal nutrition • Food of animal origin • Resource
efficiency
1 Introduction
The world population is still growing reaching seven billions in October 2011
asking for more and better food. Sustainability in feed and food production is a key
challenge for agriculture as summarized in many papers recently (Fedoroff et al.
2010; Flachowsky and Schulz 2011; Foley et al. 2011; Godfray et al. 2010; Schiere
et al. 2002) and books or proceedings (Behl et al. 2010; HPLE 2012; Welzer and
Wiegandt 2011; Wenk et al. 2009). In the future there will be a strong competition
for arable land and further non-renewable resources such as fossil carbon-sources,
water (Hoekstra and Champaign 2007; Renault and Wallender 2000; Schlink et al.
2010), some minerals (such as phosphorus; Sect. 3.2.) between feed/food, fuel,
fibre, areas for settlements and natural protected areas. According to the FAO
(2009a, b) human population will globally increase from currently about seven to
nine billion people in 2050, but the estimated need for meat and milk will nearly
double during this time (Steinfeld et al. 2006).
There is no essential need for food of animal origin as demonstrated by vega-
nians, but the consumption of meat, fish, milk and eggs may contribute significantly
to meet the human requirements in amino acids (D’Mello 2011; Pillai and Kurpad
2011; WHO et al. 2007; Young et al. 1989), some important trace nutrients such as
Ca, P, Zn, Fe, I, Se, Vitamins A, D, E, B12 etc. especially for children, juveniles as
well as for pregnant and lactating women (Wennemer et al. 2006). Human nutrition-
ists (Jackson 2007; Waterlow 1999) recommend that about one third of the daily
protein demand (0.66–1 g per kg body weight; Jackson 2007; Rand et al. 2003;
WHO et al. 2007) should originate from protein of animal origin. That implies that
about 20 g of the daily protein intake of about 60 g should base on protein of animal
origin, which is however, lower than the present average consumption all over the
world (without fish 23.9 g per day; Table 1). Table 1 shows also very large differ-
ences in protein intake and consumption of food of animal origin between “poor and
rich” countries.
In addition the conditions of production of food of animal origin are being ques-
tioned more and more, especially in the so-called developed countries as exemplified
in Fig. 1.
Immediately after the Second World War, people were hungry and required all
types of food. The so-called “Food Security” was much more important than “Food
safety” or aspects of food processing or animal health and welfare. This situation
inversed during the past years predominantly in the Western countries. Nevertheless,
Plant and Animal Breeding as Starting Points for Sustainable Agriculture 203
Table 1 Intake of milk, meat and eggs as well protein of animal origin per inhabitant and year and
portion (%) of total protein intake (Minimum and Maximum-values, global averages and Germany
for comparison; kg per inhabitant and year: data from 2005; FAO 2009a)
Food Minimum Average Maximum Germany
Milk (kg per year) 1.3 82.1 367.7 248.7
(PRa Kongo) (Sweden)
Meatb (kg per year) 3.1 41.2 142.5 83.3
(Bangladesh) (Luxembourg)
Eggs (kg per year) 0.1 9.0 20.2 11.8
(PRa Kongo) (PRa China)
Edible protein of animal origin 1.7 23.9 69.0 52.8
(g per person and day) (Burundi) (USA)
Portion of animal protein in 4.0 27.9 59.5 53.7
% of total protein intake (Burundi) (USA)
per person
a
Peoples Republic
b
Probably empty body weight (meat plus bones)
Question/Task
I‘m hungry! I’d like something I‘m nervous! How can we feed
Consumer Is there to eat! How safe the world ?
anything to eat? What do we have? is my food?
Securing the food supply, Food Quality, Food Safety Field of conflicts
Manufacturing enough food Reducing surpluses between feed, food
and fuel
Policy
„ Food Security “ „Food Safety“ Food Security and

Food Safety
Increase in agricultural Quality research, Research on Effective use of

production Product quality safety by-products;
Agricultural GM-crops
Research Use of all resources Process quality
Effective (conservational) Life Cycle
use of resources Studies
1945 1950 1960 1970 1980 1990 2000 2010 2020

Year
Fig. 1 Dominating questions for food production as well challenges for policy and agricultural
research after the Second World War in Europe (After Flachowsky 2002a). GM: genetically modified
the fundamental question “Is there anything to eat?” (see Table 1 and Fig. 1) is still
relevant for about one billion people (HPLE 2012; WHO et al. 2007) in many coun-
tries which clearly demonstrate the necessity of producing more and better food of
plant and animal origin all over the world.
On the other hand there is a high variation in the availability and consumption of
food of animal origin between individuals and in countries between 1.7 and about
70 g protein of animal origin per person and day (see Table 1). Other reasons for
Table 2 Effects of animal species, categories and performances on some emissions (per kg edible
protein, Flachowsky 2002b, 2011; see Table 4 for fundamentals of calculations)
Nitrogen Methane Emissions in kg per kg edible

Protein source Performance excretion emission protein
(Body weight) per daya (% of intake) (g per day)b P N CH4b CO2eqc
Dairy cow 10 kg milk 75 310 0.10 0.65 1.0 30
(650 kg) 20 kg milk 70 380 0.06 0.44 0.6 16
40 kg milk 65 520 0.04 0.24 0.4 12
Dairy goat 2 kg milk 75 50 0.08 0.5 0.8 20
(60 kg) 5 kg milk 65 60 0.04 0.2 0.4 10
Beef cattle 500 gd 90 170 0.30 2.3 3.5 110
(350 kg) 1,000 gd 84 175 0.18 1.3 1.7 55
1,500 gd 80 180 0.14 1.0 1.2 35
Growing/ 500 gd 85 5 0.20 1.0 0.12 16
fattening 700 gd 80 5 0.12 0.7 0.08 12
pig (80 kg) 900 gd 75 5 0.09 0.55 0.05 10
Broilers (1.5 kg) 40 gd 70 Traces 0.04 0.35 0.01 4
60 gd 60 0.03 0.25 0.01 3
Laying hen 50%e 80 Traces 0.12 0.6 0.03 7
(1.8 kg) 70%e 65 0.07 0.4 0.02 5
90%e 55 0.05 0.3 0.02 3
a
See Table 4 for fundamentals of calculation
b
CH4-emission depending on composition of diet
c
Adequate to Carbon Footprints (Sum of greenhouse gas emission of CO2; CH4 (× 23) and N20
× 300; IPCC 2006) for edible protein of animal origin
d
Body weight gain
e
Laying performance
foods of animal origin are the high bioavailability of nutrients and their considerable
enjoyment value. Such food is also considered as an indicator for the standard of
living in many regions of the world. Other reasons for the higher demand of food of
animal origin in some countries are the increased income of the population (Keyzer
et al. 2005) and the imitation of the so-called “Western style of life” (nutrition).
Higher food amounts of animal origin require higher plant yields and/or more area
for feed production and more animals and/or higher animal yields.
In addition feed/food production causes emissions with a certain greenhouse
gas potential such as carbon dioxide (CO2) from fossil fuel, methane (CH4; green-
house gas factor (GHF) about 23; IPCC 2006) from the enteric fermentation esp.
in ruminants and from the excrement management as well as nitrogen compounds
(NH3; N2O: GHF about 300; IPCC 2006) from the protein metabolism in the ani-
mals (DEFRA 2006; FAO 2010; Flachowsky et al. 2011; Flachowsky and
Hachenberg 2009; Godfray et al. 2010; Grünberg et al. 2010; Leip et al. 2010; see
Table 2). Apart from the low input of limited resources along the food chain (Fig. 2)
a low output of minerals such as phosphorus and some trace elements and green-
house gases (CO2, CH4 and N2O) during feed/food production are very important
aims of sustainable agriculture. Table 2 shows significant differences in emissions
Fertilizer, Fuel, Fuel, Electricity, Fuel,

Fuel, Electricity Electricity Fuel Electricity
Inputs Herbicids,
Seeds,
Water
1)
(CO2) Food of animal
Men
origin
Elements of Soil, Feed,
food chain Harvest, Mixed feeds, Animals
Plant
Storage By-products Biogas,
Excrements Manure
Outputs of
1)
greenhouse CO2 N2O CO2 CO2 CH4 (CO2) CH4 N2O CO2 CH4
gases
1) CO2 will be fixed by photosynthesis and produced by animal metabolism, therefore

it is considered as emissions-neutral
Fig. 2 Substantial elements of the chain to produce food of animal origin as well as selected
inputs of limited resources and outputs of greenhouse gases (Flachowsky and Hachenberg 2009)
between animal species/categories as well as animal yields per kg edible protein.

Such aspects should be more considered in future assessments about food security,
resource efficiency and emissions.
Based on the statement before sustainable agriculture may be defined as agricul-
ture with (1) low input of limited natural resources such as water, arable land, fuel
and other substances, (2) a very efficient use of such resources to satisfy human
food needs and other products (3) and low emissions/pollution to enhance environ-
mental quality and and to protect natural resources (Fig. 2).
Therefore the objective of the present paper is to evaluate the potentials of plant
and animal breeding for the most efficient use of non-renewable resources and their
consequences for emissions to contribute to sustainable agriculture.
2 Challenges for Plant Breeding
Plant breeding and cultivation are the key elements and starting points for feed and
food security during the next years (Flachowsky 2008; SCAR 2008; The Royal
Society 2009). The most important objectives for plant breeders can be summarized
as follows:
– High and stable yields with low external inputs of non-renewable resources (low
input varieties) such as water, minerals, fossil fuel, plant protecting agents etc.
(see Table 3)
– Maximal use of naturally unlimited resources such as sunlight, nitrogen and car-
bon dioxide from the air (see Table 3)
– Higher resistance against biotic and abiotic stressors
– Stable animal health and adaptation to potential climate changes
– Optimisation of the genetic potential of plants for a highly efficient photosynthesis
– Lower concentrations of toxic substances such as secondary plant ingredients,
mycotoxins from toxin-producing fungi, contaminants from anthropogenic
activities or geogenic origin
Table 3 Potentials to produce phytogenic biomass and

their availability per inhabitant under consideration of
the increase of population (Flachowsky 2010; The Royal
Society 2009)
Plant nutrients in the air (N2, CO2) ↑↔
Solar energy ↔
Agricultural area ↓
Water ↓
Fossil Energy ↓
Mineral plant nutrients ↓
Using of potentials of the genetic pool ↑
↑ Increase, ↓ Decrease, ↔ No important influence
– Lower concentrations of substances that influence the utilisation or bioavailability

of nutrients such as lignin, phytate, enzyme inhibitors and tannins
– Higher concentrations of the nutritive value determining components such as
nutrient precursors, nutrients, enzymes, pro- and prebiotics and essential oils.
From the global perspective of feed and food security, low input varieties with
high and stable yields should have the highest priority in plant breeding. In addition,
low losses on the field, during harvest and storage are also important aspects of
feed/food security. Furthermore, undesirable substances can often not be removed
from feedstuffs or can only be removed with great effort (Fink-Gremmels 2012;
Flachowsky 2006; Verstraete 2011). Therefore a decrease of undesirable sub-
stances in plants is also an important objective of plant breeding. From the perspec-
tive of nutrition, an increase of essential nutrients such as amino acids, fatty acids,
trace elements, vitamins etc. (Beauregard and Hefford 2006; Bouis 2002; Napier
2007; Lucas et al. 2007; Ufaz and Galili 2008; Welch and Graham 2004; White and
Broadley 2005), could be very favourable to meet the requirements of humans and
animals for essential nutrients although this is less important in some parts of the
world like Europe due to the availability of large amounts of food and feed additives
on the market. It could be also important to create food/feed qualities as proteins
with bioactive peptides, which may contribute to improve human and animal health
(Maruyama et al. 2011). Furthermore, potential aspects of climate changes (HPLE
2012; IPCC 2012) should be considered by plant breeders and “new” plants should
be adapted to such changes (Newman et al. 2011; Reynolds 2010). It is possible to
fulfil the objectives of plant breeding mentioned above with conventional breeding
(Flachowsky 2012) however in the future methods of “green” biotechnology may be
more flexible, more potent and faster (Whitford et al. 2010). “New” plants, newly
expressed proteins in plants and/or changed composition of plants are real challenges
for animal and human nutritionists for safety and nutritional assessment of such
products (see Fig. 4).
Increasing feed/food demands require higher plant yields and/or more area for
production (see Table 13). Because of some limited resources, low input plants are
an important prerequisite for solving future problems and establishing a sustainable
agriculture. Such plants should be very efficient in the use of mineral plant nutrients
(incl. N), fuel, water and arable land (high yields), but they should as well be able to
use sun energy and unlimited plant nutrients from the air such as N2 and CO2 (see
Table 3). Non legumes should also be able to use N from the air for N-fixing symbio-
sis. Furthermore the genetic pool available in plants, animals and microorganisms
should contribute to optimizing plants and animals for a more efficient conversion of
limited resources into feed and food. Maintaining the biodiversity of the available
genetic pool is also a very important aspect of sustainable agriculture.
Subsequently animal feeding studies are necessary to demonstrate the digest-
ibility/availability of the changed composition of the plants or the newly, or in higher
amounts, expressed nutrients (see Fig. 4 and Sect. 4 for some examples).
Expected climate change may become an additional challenge for plant breeders.
Some authors (Easterling et al. 2007; Reynolds 2010) predict a 15–20% fall in
global agriculture production by 2080 in the consequence of the following expected
climate change (Whitford et al. 2010):
– Adaption to greater extremes in climate conditions
– Limited or more variable water supply
– Increasing soil salination
– Higher disease infection and pest infestations.
A rapidly changing climate will require rapid development of new plant variet-
ies. The negative effects of climate change could be greater than the possible solu-
tions by conventional plant breeding. Therefore a large “technology gap” between
solutions by conventional breeding and need for adaptation to climate change
will result in adequate or lower plant yields (Whitford et al. 2010). Plants undergo
adaptive change as they acclimatize to new environments. Drought-resistance, high
water-use-efficiency, heat tolerance and disease resistant plants would be important
objectives of plant breeding under climate change, combined with techniques
enhancing speed, flexibility and efficiency of plant breeding triggering the “Second
Green Revolution”.
3 Challenges for Animal Breeding
Domestic animals are very important for most people and are of special interest for
farmers all over the world. They produce well tasting and valuable food of animal
origin, but they also supply mankind with power, fertilizer, clothing, heating mate-
rial and many other things including security for farmers and their families, insur-
ance and prestige. Conversion of feed into food of animal origin is associated with
high energy and nutrient losses and environmental pollution, including excretion
of Nitrogen (N), Phosphorus (P), trace elements, Methane (CH4) etc. (see Table 2)
The resources such as fuel, water, arable land and specific minerals such as P are
limited (see Table 3), but needed for synthesis of phytogenic biomass as fodder for
animals (Flachowsky 2008). Only a small proportion of feed is converted into food
products by the animals. This amount is variable and depends on animal species,
the performance of the individual animal and several other factors. Most of the
substances are emitted into the environment (Table 2).
The main purpose for animal husbandry in great quantities is the production of
valuable protein in the form of milk, eggs, meat and/or fish for human consumption.
Large differences exist in protein yields per animal or per kg body weight depending
on species and production category as well as performance status (see Table 2).
Animal husbandry is considered as an important source of greenhouse gas emis-
sions because of the high global warming potential of their emissions (see Table 2).
So-called Carbon Footprints (CF; Live Cycle Assessments (LCA), Eco-Balances)
consider the greenhouse gas potentials of climate relevant gases and are given in
CO2eq per g or per kg product. The objectives of CF are to sensitize producers and
consumers for an efficient use of fossil carbon sources and to reduce greenhouse gas
emissions (GHG) per unit of product. Various authors calculated the contribution of
animal husbandry between about 10% (Lesschen et al. 2011; O’Mara 2011) and up
to 20% (Steinfeld et al. 2006) of global greenhouse gas emissions. Herrero et al.
(2011) analysed some references and found a range between 8% and 51% of GHG
from livestock.
Ruminants contribute substantially to global methane emission (de Vries and de
Boer 2010; Flachowsky and Brade; 2007; Janssen 2010; Jouany 2008; Kebreab
et al. 2006). Carbon Footprints per unit of product are indicative for CO2 emissions
and serve as an important indicator for determining the efficiency of food produc-
tion. In the case of non-ruminants there is a nearly linear correlation between fossil
energy input and emission measured as CF, but in ruminants between 50% and
80% of CF are based on methane emissions (Fig. 3). Therefore it is difficult to
compare CF by ruminants and non-ruminants and to make conclusions on the fuel
input. A more thought-out and detailed assessment of this subject has recently
been made by Flachowsky and Kamphues (2012).
The FAO (2010) calculated Carbon Footprints on a global scale and found values
between 1.3 kg CO2eq per kg milk in North America and Europe and 7.5 kg CO2eq.
per kg milk in sub-Saharan Africa. Selected emissions with their corresponding
CO2-footprints per kg edible protein are shown in Table 2. To reduce greenhouse gas
emission of the global animal population, the ratio of production/consumption of
feed of animal origin has to be reduced and the production of phytogenic biomass
has to be increased. The overall efficiency of feed conversion into animal derived
food for human consumption has to be improved and should be one of the most
important challenges for animal breeders. The breeding of domestic animals has a
longstanding history, starting with domestication several 1,000 years ago. Using the
technical options that were available in each time period, humans have propagated
those populations that deemed useful for their respective needs and purposes.
Nowadays genomics offers great opportunities to improve livestock production all
over the world, resulting in a so-called “livestock revolution” (Laible 2009; Niemann
et al. 2010; Rothschild and Plastow 2008). Presently, the perspectives for a feed-
efficient production of animal derived food for human consumption with significantly
reduced emission of greenhouse gases are the main objectives of animal breeding.
CO2Eq (kg per kg edible Protein)

120
100
80
Methane-Portion
60
40
20
0
Performance 10 20 40 2 5 500 1000 1500 500 700 900 40 60 50 70 90
per day kg Milk kg Milk g BWG g BWG g BWG % LP
Animal Dairy cow Dairy goat Beef cattle Growing pig Broiler Laying hen
species/category
Fig. 3 Carbon Footprints (kg CO2eq) per kg edible protein of animal origin in dependence of animal
species/-category and animal performance under consideration of the methane emission (BWG
body weight gain, LP laying performance; Flachowsky et al. 2011)
Strategies to produce such animals were recently summarized by Niemann et al.

(2011) and include:
• Higher feed intake by animals (Forbes 1995; Langhans 2010; Schwartz et al.
2000) to improve the ratio between energy/nutrient requirements for mainte-
nance and animal yields (Tables 4 and 5) and to reduce animal numbers
• Higher digestibility of feed (Table 6) and higher absorption of digested nutrients
to make energy/nutrients more available from the same feed amount
• Lower energy/nutrient requirements for maintenance of the animals (Table 7);
reduction of energy losses in the digestive tract (e.g., CH4, Table 8)
• Lower energy need for protein synthesis in the body or increase of anabolic
processes and lower catabolic processes in the animal
• Lower fat content in animal bodies or lower excretion of fat in milk and eggs or
lower excretion of lactose in milk (lower energy content in products)
• Improved animal health, specifically animals with higher resistance against biotic
and/or abiotic stressors (incl. climate changes; Laible 2009) and lower losses
during production may also contribute to a more efficient conversion of feed.
The Tables 4, 5, 6, 7, and 8 show some examples for more efficient feed conversion
and reduction of emissions by animals. The amount of feed intake depends on animal
species and production categories, physiological stage, body weight, feed quality and
structure and many other factors as shown in Table 4. Furthermore the roughage-to-
concentrate ratio of diets, edible fraction and protein content of edible fractions, amounts
of edible protein per day and per kg of body weight of animals are shown in Table 4. The
210
Table 4 Effects of animal species, production categories and performance on yields of edible protein per animal or per kg body weight and day (Flachowsky 2002b)
Roughage
Dry matter to concen-trate Edible fraction Protein in edible Edible protein
Protein source Performance intake ratio (on dry (% of product fraction (g per Edible protein (g per kg body
(Body weight) per day (kg per day) matter base, %) or body mass) kg fresh matter) (g per day) weight)
Dairy cow 10 kg milk 12 90/10 95 34 323 0.7
(650 kg) 20 kg milk 16 75/25 646 1.0
40 kg milk 25 50/50 1,292 2.0
Dairy goat 2 kg milk 2.0 80/20 95 36 68 1.1
(60 kg) 5 kg milk 2.5 50/50 170 2.8
Beef cattle 500 ga 6.5 95/5 50 190 48 0.14
(350 kg) 1,000 ga 7.0 85/15 95 0.27
1,500 ga 7.5 70/30 143 0.41
Growing/ 500 ga 1.8 20/80 60 150 45 0.56
fattening 700 ga 2.0 10/90 63 0.8
pig (80 kg) 1,000 ga 2.2 0/100 81 1.0
Broiler 40 ga 0.07 10/90 60 200 4.8 3.2
(1.5 kg) 60 ga 0.08 0/100 7.2 4.8
Laying hen 50%b 0.10 20/80 95 120 3.4 1.9
(1.8 kg) 70%b 0.11 10/90 4.8 2.7
90%b 0.12 0/100 6.2 3.4
a
Body weight gain
b
Laying performance
G. Flachowsky et al.
Table 5 Model calculation to show the influence of dry matter intake (Dry matter intake: 7.0 MJ
Net energy lactation/kg DM) of dairy cows (body weight: 650 kg; 4% milk fat; GfE 2001) on
energy intake, percentage of maintenance, milk yield, energy per kg of milk as well as emissions
per kg of milk (Niemann et al. 2011)
Dry Matter Intake (DMI, kg per day) 10 15 20 25 30
Energy intake (MJ NEL per day) 70 105 140 175 210
Maintenance (37.7 MJ NEL per cow 53.9 35.9 26.9 21.5 18.0
per day; % of total NEL-Intake)
Milk yield (3.3 MJ NEL per kg) 9.8 20.4 31.0 41.6 52.2
Net energy per kg milk (MJ NEL 7.1 5.1 4.5 4.2 4.0
per kg milk)
Methane emissiona
(g per day) 240 360 480 600 720
(g per kg milk) 24.5 17.6 15.5 14.4 13.8
Carbon footprint (g CO2eq per kg milk)b 825 605 530 495 475
a
According to Flachowsky and Brade (2007): 24 g CH4 per kg DMI for all diets
b
Calculated on the base of the greenhouse potential of CH4 (× 23) and the calculations by Daemmgen
and Haenel (2008)
Table 6 P-digestibility (%) of non-transgenic and transgenic pigs (enviro-pigs) using soybean
meal as the sole source of P (Golovan et al. 2001b)
Pigs (Body weight) Non-transgenic Transgenic (Environment pig)
Weaning (6–15 kg) 48.5 ± 5.4 87.9 ± 3.4
(n = 16) (n = 14)
Growing-finishing (20–65 kg) 51.9 ± 10.3 98.8 ± 3.4
(n = 16) (n = 14)
Table 7 Model calculation to show the influence of various energy maintenance requirements on
milk yield of lactating cows (Body weight: 650 kg per cow, Dry matter intake: 20 kg per day; Net
energy content of feed: 7.0 MJ NEL per kg DM; Niemann et al. 2011)
Maintenance requirements for energy
(MJ NEL per kg BW0.75) 0.2 0.25 0.3 0.35 0.40
(MJ NEL per cow per day) 25.7 32.2 38.6 45.0 51.5
Energy intake (MJ NEL per cow per day) 140 140 140 140 140
Maintenance in % of NEL-intake 18.4 23.0 27.6 32.1 36.8
Milk yield (kg per cow per day) 34.6 32.7 30.7 28.8 26.8
Methane emission
(g per cow per day) 480 480 480 480 480
(g per kg milk) 13.9 14.7 15.6 16.7 17.9
Carbon footprints (g CO2eq per kg milk) 480 505 535 575 615
Table 8 Model calculation to show the influence of methane reduction on the energy available for
dairy cows and milk yields (Conditions for calculation: Dry matter intake: 20 kg per cow per day;
Body weight: 650 kg per cow, 7 MJ NEL per kg DM with 20 g CH4-emission; Niemann et al. 2011)
Methane production
(g per kg DMI) 30 25 20 15
(g per cow per day) 600 500 400 300
Energy intake (MJ NEL per day) 130 135 140 145
Milk yield (kg per day) 28.0 29.5 31.0 32.5
Methane emission (g per kg milk) 21.4 17.0 12.9 9.2
Carbon footprint (g CO2eq per kg milk) 735 585 440 315
data of Table 4 are also the basis for calculations shown in Table 2. Examples for a more
efficient feed conversion are described in Sects. 3.1, 3.2, 3.3, and 3.4.
3.1 Higher Feed Intake
Increased feed intake may contribute to a higher energy and nutrient intake and thus
improve the ratio between energy available for performance and for maintenance as
demonstrated for dairy cows (Table 5).
Higher milk yield resulted in higher methane emission per cow, but significantly
lower methane and Carbon Footprints per kg of milk (Table 5). On the other hand,
the higher milk yields result in less dairy cows and beef cows are required to produce
a certain amount of beef. So-called allocations (Cederberg and Stadig 2003; Feitz
et al. 2007; Flysjö et al. 2011; Thomassen et al. 2008; Zehetmeier et al. 2011) are
necessary to assess CF for milk and beef production. For example, Zehetmeier et al.
(2011) made such an economic allocation of GHG-emissions in dairy husbandry
(milk and beef) on the base of 6,000; 8,000 or 10,000 kg milk per cow and year and
asked for the same amount of beef. The total GHG-emission for milk production on
farm decreased from 1.06, 0.93 to 0.89 kg CO2eq/kg milk; those of beef increased
from 10.75; 13.13 to 16.24 kg CO2eq/kg beef. Such and similar calculations (e.g., on
the basis of 1 kg edible protein) are recommended to assess emissions for milk/meat
of various production intensities (see Flachowsky and Kamphues 2012).
3.2 Higher Digestibility
Unfortunately, in ruminants the strategy of higher feed intake may reduce the digest-
ibility of organic matter to a certain degree (Gabel et al. 2003; NRC 2001). This
decrease in digestibility of feed in the gastrointestinal tract of animals could be
overcome by specific plant breeding programmes (Sect. 2.; Stein and Rodriguez-
Cerezo 2009) or a higher expression of specific digestive enzymes by microorganisms
in the rumen or in other parts of the digestive tract.
In non-ruminants there are some examples that genetically modified animals are
able to express enzymes, e.g. phytase in higher amounts (Golovan et al. 2001a) in
mice; Golovan et al. (2001b) in pigs and Cho et al. (2006) in poultry. Golovan et al.
(2001b) used the parotid secretary protein promoter linked to the E. coli appA
phytase gene and produced transgenic piglets which were able to produce phytase
and to increase P-digestibility from plants (Table 6). The authors consider such
developments as a contribution to help ameliorating the planets “Phosphate crisis”
(Cho et al. 2006). The transgenic enviropigs express E. coli phytase and showed
also after some generations the introduction of a novel transgene into the animal
genome (Golovan et al. 2008). Further research incl. food safety of products of such
animals is necessary to establish those animals on farm level. In April 2012 the
financial support for the Enviropig programm was stoped and in June 2012 all pigs
from the 10th generation were killed (Leung 2012; Schimdt 2012). However, the
genetic material was stored at the Canadian Agricultural Genetics Repository
Program.
3.3 Lower Requirements for Maintenance
Another way for a more efficient conversion of feed into food of animal origin could
be a lower need of energy and nutrients for maintaining the animals. These require-
ments depend on animal species, production category, body composition and other
factors. The energy requirements are usually given per kg metabolic body size (kg
BW0.75; e.g., 0.293 MJ NEL/kg BW0.75 for dairy cows; GfE 2001). Similar values
were deduced by other energy evaluation systems (e.g., 0.272 or 0.335 MJ NEL/kg
BW0.75 for beef or dairy cattle by the NRC 2001). All the values are characterized by
a large range between individuals. Lower maintenance requirements save energy/
nutrients and more energy/nutrients are available for animal yield with consequences
for lower methane emission and Carbon Footprints per unit product (Table 7).
3.4 Lower Methane Emissions
During anaerobic microbial fermentation some feed energy is lost via methane
(Baldwin 1995; Kebreab et al. 2006; 2008; Martin et al. 2010). The methane
losses vary between 4% and 10% of the gross energy intake of ruminants
(Flachowsky and Brade 2007; Tamminga et al. 2007) and increase with increasing
fibre content of the rations (Ellis et al. 2010; Hindrichsen et al. 2005; Kirchgessner
et al. 1995). The consequences of lower methane emission on the yield and emis-
sions of dairy cows are shown in Table 8 and described in many papers on feeding
(Beauchemin et al. 2008; Fievez et al. 2010; Flachowsky and Brade 2007;
Flachowsky et al. 2011; Kreuzer 2009; for more details see: Greenhouse Gas in
Animal Agriculture – Finding a Balance between Food and Emissions. Anim.
Feed Sci. Technol. 2011; 166–167; 1–796). Some breeding measurements (Beever
et al. 2001; Brade et al. 2008; Hegarty 2004; Hegarty et al. 2007) may also
influence the methane emissions by ruminants.
Apart from those aspects mentioned above, there are some other opportunities
for a more efficient conversion of feed such as
– Lower fat content in the food of animal origin (e.g., meat, milk, eggs)
– Higher protein content in the products (e.g., milk)
– Lower lactose in milk to relieve metabolism (liver) by lower gluconeogenesis
Further details of the current stage for a more sustainable animal production are
described by Laible (2009) and Niemann et al. (2011).
4 Animal Nutrition Between Plant and Animal Breeding
Animal nutrition has to contribute and to act as an important element in the food
chain between plant breeding and plant production on one hand and animal breeding
and animal husbandry on the other hand (Fig. 4). In this function animal nutritionists
may substantially contribute to a sustainable production of food of animal origin.
First of all they have to analyse the composition of plants and to assess the digest-
ibility/bioavailability of changes in plants (lower content of undesirable substances
such as lignin, phytate, etc., or higher content of valuable ingredients such as amino
acids, fatty acids, minerals, vitamins or nutrient precursors), but also in plants with
improved use of limited natural resources or better conversion of CO2 as plant nutri-
ent (DaMatta et al. 2009; Lohoelter et al. 2012; Porteaus et al. 2009). Animal feed-
ing studies to assess the changes in plants should be done according recommendations
given by EFSA (2008); Flachowsky and Böhme (2005) or ILSI (2007). Apart from
bioconversion and assessment of sustainability such studies should also demon-
strate the influence of changed plants/feed on animal health and performance as
well as composition, the quality of food of animal origin and the transfer of specific
nutrients into food of animal origin. The following paragraphs show examples of
some changes in plants and the challenges for animal nutritionists for nutritional
assessment of changes (Tables 9, 10, and 11).
4.1 Higher Concentration of a Nutrient Precursor (ß-Carotene)
ß-carotene is present in many plants and it is the most important precursor for vita-
min A in humans and animals. Table 9 shows a study to assess the bioconversion of
ß-carotene into vitamin A in a model animal Mongolian gerbils. Different diets
Animal nutrition Animal breeding

Plant breeding (Nutritional and safety (inl. Clones and GM-
(inl. GMO) assessment of feed) animal)
Fig. 4 Animal nutrition and nutritional assessment of feeds between plant and animal breeding
Table 9 Experimental design to assess the conversion of various sources of ß-carotene into vitamin
A and to compare it with vitamin A storing in the liver in Mongolian gerbils (60% maize in diets;
n = 10, depletion phase: 4 weeks, feeding: 8 weeks; Howe and Tanumihardjo 2006)
Unsupplemented
control (Maize Control + Control +
poor in carotene) Carotene rich maize β-carotene vitamin A
β – carotene 0 8.8 8.8 4.4
(nmol/g)
Theoretical retinol 0 106 106 106
intake (nmol/d)
Retinol in serum 1.23 1.25 1.23 1.22
(μmol/l) ±0.20 ±0.22 ±0.20 ±0.16
Retinol in liver 0.10a 0.25a 0.25a 0.56a
(μmol/g) ±0.04 ±0.15 ±0.08 ±0.15
a
Means with different letters differ significantly (p < 0.05)
Table 10 Concentrations of some n-3 fatty acids (mg/100 g fresh tissue) in body
samples of broilers (Rymer et al. 2011)
+45 (grower) and 50 g SDA-oil
Sample Control (finisher diet) per kg feed
Breast meat
C18:4n-3 3 231
C20:5n-3 12 28
C22:6n-3 7 14
Leg meat
C18:4n-3 10 442
C20:5n-3 5 53
C22:6n-3 8 21
Skin
C18:4n-3 111 3,673
C20:5n-3 31 317
C22:6n-3 21 78
were fed after a depletion period. The results show that the retinol concentration of
liver of gerbils fed with carotene-rich maize was similar to those animals fed with
maize poor in carotene and supplemented with adequate amounts of ß-carotene.
That means in this case that ß-carotene from maize is almost identically converted
into vitamin A as supplemented ß-carotene, but it is stored in the liver to a lower
extent than vitamin A.
In the case of “Golden Rice” containing between 1.6 (Ye et al. 2000) and up to
37 mg ß-carotene per kg dry rice (Paine et al. 2005), the first studies to determine
the vitamin A value of ß-carotene were done with deuterium-labelled rice in five
humans (Tang et al. 2009) and not in laboratory or target animals.
Results demonstrate that plant breeding is able to introduce genes in plants which
express nutrients in higher amounts with adequate bioavailability to supplemented
ß-carotene.
Table 11 Conventional and low-phytate maize (78.5% of the mixture) in the feed of fattening pigs
(from Spencer et al. 2000)
Groups Control Low-phytate maize
(0.3 g of available (1.7 g of available
Parameters P per kg) P per kg)
Inorganic P supplement − + − +
P content (g/kg)
29–73 kg live weight 3.4 5.4a 3.4 5.4a
73–112 kg live weight 3.2 4.7 3.2 4.7b
Feed intake (kg/d) 2.23c 2.50c 2.53c 2.51c
Live weight gain (g/day) 730 870c 900c 880c
Feed per gain (kg/kg) 3.05c 2.87c 2.81c 2.85c
P excreted (g/kg weight gain) 4.6c 8.9c 3.8c 8.8c
Strength (4th metacarpal bone, kg) 79.3c 138.5c 132.2c 153.9c
Ash content (% in 4th metacarpal bone) 53.5c 60.1c 59.3c 61.2c
a
+2.0 g P/kg
b
+1.5
c
Different letters in one line indicate significant differences (p < 0.05)
18:0 Δ9 18:1 Δ12 18:2 Δ15 18:3

Stearic acid Oleic acid Linoleic acid a -linolenic acid
LA ALA
Δ6 Δ6
18:3 Δ15 18:4

g -linolenic acid Stearidonic acid
GLA SDA
Fig. 5 Fatty acid biosynthesis in plants and the new introduced changes to produce Stearidonic acid
(C18:4 n-3) and the effects of various desaturases (Δ6 and Δ15; by Ursin 2003 and Whelan 2009)
4.2 Expression and Using of Stearidonic Acid in Soybeans

(C18:4 n-3; SDA)
Plant oils can provide renewable sources of high value-fatty acids (Dyer et al. 2008;
Lunn and Theobald 2006). For example, the introduction of two new genes in
soybeans effecting the expression of Δ6- and higher expression of Δ15-desaturases
causes the biosynthesis of a highly unsaturated fatty acid with four double bonds
(Stearidonic acid; see Fig. 5).
Stearidonic soybean oil contains between 20% and 30% Stearidonic acid (SDA;
C18:4 n-3). Rymer et al. (2011) added 45 (grower) and 50 g SDA-oil containing 24%
of Stearidonic acid per kg finisher broiler diet and compared this with conventional
soybean oil. The authors did not observe any significant influence of SDA-oil on feed
intake, weight gain and feed conversion rate in the animals, but they found higher
concentrations of Stearidonic acid as well as C20 and C22 polyunsaturated fatty acids
in various body fats (Table 10). Similar results are described by Bernal-Santos et al.
(2010) in lactating cows after duodenal infusion of SDA-soybean oil, by Kitessa and
Young (2011) after feeding of ruminal protected SDA-oil to dairy cows and by Meja
et al. (2010) in laying hens.
Such fatty acids could have health effects (Lunn and Theobald 2006) and may be
transferred in long-chain polyunsaturated fatty acids with 20 and 22 C-atoms (see
Table 10) and would be able to replace fish oils in human and animal nutrition.
4.3 Lower Phytate in Maize and P-Utilization in Pigs
Phytate is one of the most important inhibitors of phosphorus-bioavailability. The

reduction of the undesirable substance phytate (Raboy 2002) or the expression of
the enzyme phytase (Chen et al. 2008; Gao et al. 2012) in grains or in animals (see
Table 6) as well as the supplementation of phytase as feed additive to diets of non-
ruminants increases the P-availability in non-ruminants as shown in Table 11 for
low phytate maize in pigs. The unsupplemented low-phytate maize resulted in
nearly the same body weight gain of pigs (P > 0.05) as the control maize supple-
mented with mineral P, but in much lower P-excretions per kg body weight gain
(P < 0.05) and therefore in a more resource efficient and sustainable production.
4.4 Consequences for Animal Nutritionists
Progress in animal breeding may also have consequences in energy and nutrient
requirements of food producing animals. The energy and nutrient requirements of
newly bred animals must be re-evaluated after certain intervals. For example,
national scientific committees, such as the National Research Council (NRC) of the
USA or the Society of Nutrition Physiology (GfE) in Germany update their require-
ments for dairy cows (e.g., NRC; since 1944 7th ed.), pigs (NRC: since 1944 10th
ed.) or poultry (NRC: since 1944 9th ed.) every 5 to 15 years. Under consideration
of spontaneous breeding progresses (e.g., higher expression of specific enzymes or
changes in the composition of animal products; see Tables 5, 6, 7, and 8) specific
recommendations for energy and the nutrient supply of animals could be given
between dates of general revision.
5 Conclusion
Plant and animal breeding are the starting points of sustainable resource usage along
the food chain. Table 12 shows objectives of plant breeding, their present significance
in practise and the contributions to food safety and food security as well as for the
sustainability of food production. High and stable yields and a more efficient use of
non renewable resources should be the main aim of plant breeding and contribute
Table 12 Assessment of present modifications of plants from the view of food safety, food security
and sustainability of food production
Contributions to
Present Food Global Sustainability
Objectives of plant breeding significance safety food security of food production
More tolerant against herbicides ↑↑↑ ↑↑ ↑ (↑)
More resistant against insects etc. ↑↑ ↑ ↑ ↑
More valuable ingredients ↑ ~ (↑) ↑
Less undesirable ingredients (↑) ↑↑ ↑ ↑↑
More efficient use of non-renewable (↑) ↑ ↑↑↑ ↑↑↑
resources (water. nutrients,
area etc.)
↑↑↑ Extremely high, ↑↑ Very high, ↑ High, ~ Not important
Table 13 Model calculation on the influence of human intake of protein of animal origin (except
fish), yields of plants and performances of animals as well relation between protein from meat and
milk on need for arable area (see papers by Flachowsky and Bergmann 1995; Flachowsky 2002b
and Flachowsky et al. 2008 based on other plant yield levels and animal performances)
Protein of animal origin (g per inhabitant per day)
Relation between 10 20 40 60
meata and milk Intensity level
(% of protein) Ab Bc A B A B A B
Arable land required for feed production (mc per inhabitant per year)d
70 : 30 345 95 690 190 1 380 380 2 080 570
50 : 50 290 85 580 170 1 160 340 1 740 510
30 : 70 235 75 470 150 940 300 1 410 450
a
Relation between beef : pork : poultry meat = 20:50:30
b
Plant yield level A: 3 t DM (dry matter) cereals; 10 t DM roughage per ha and year; Performance
of animals A (per animal per day): 15 kg milk; weight gain: beef: 600 g; pork: 400 g; poultry: 30 g
c
Plant yield level B: 8 t DM cereals; 20 t DM roughage per ha and year; Performance of animals
B: 30 kg milk; weight gain: beef: 1 200 g; pork: 800 g; poultry: 60 g
d
See Tables 2 and 4 for some details of calculations (e.g., Dry matter intake; concentrate : roughage
ratio; yields of edible protein)
substantially to global food security and sustainability of food production (Table 12;
see Potrykus and Ammann 2010). The conversion of feed into food of animal origin
is associated with high energy and nutrient losses and emissions in the environment.
Therefore animal breeders may also substantially contribute to a more efficient
conversion of feed. More public supported research in these fields seems to be
necessary to reach this objective.
Higher plant yields and more efficient conversion of feed into food of animal origin
may also contribute to a lower need for arable land per inhabitant and are important
challenges for all those working in the field. Apart from plant yields and animal
performances (see Tables 2 and 4) the need for area depends also on the amount of
protein intake and the proportion between milk and meat in human nutrition as
shown in Table 13. Such calculations to assess the human need for arable land and
further non renewable resources may contribute to show possibilities and limitations
along the food chain. Furthermore, it allows drawing conclusions for a sustainable
production for sufficient food of animal origin.
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Medicinal Plant Active Compounds Produced
by UV-B Exposure
Rima Kumari and Majeti Narasimha Vara Prasad
Abstract Ultraviolet (UV) radiation is a part of the sunlight reaching Earth surface.
The UV spectrum of solar radiation is by convention divided into three parts: UV-A:
310–400 nm, UV-B: 280–310 nm and UV-C: less than 280 nm. UV-B is the most
energetic component reaching Earth surface because the stratospheric ozone layer
effectively absorbs completely wavelengths below 290 nm. UV-B is an increasing
threat due to increasing UV-B levels on Earth surface as a consequence of depletion
of stratospheric O3. In general, the effects of atmospheric UV-B radiation are nega-
tive for biological life. Enhanced levels of UV-B radiation can indeed negatively
change plant physiological processes, growth and productivity. However, while
studying UV-B effects on medicinal plants, some interesting phenomena have been
discovered. For example, basil plants respond positively to UV-B radiation by
increasing oil yield (Chang et al. J Horticult For 1:27–31, 2009). In other studies
medicinal plants show beneficial aspects in term of increase in volatile oil yield
and secondary metabolite production (Kumari et al. Ecotoxicol Environ Safety
72:2013–2019, 2009c, 2010). Medicinal herbs have great market value in India and
worldwide. The medicinal value of plants depends upon phenolics, antioxidants and
volatile yield. Therefore further UV-B experiments increasing the levels of these
compounds are needed.
Here we review the effect of UV-B exposure on metabolites, volatiles, and anti-
oxidants potential in medicinal plants. This chapter reports: (1) aspects of the global
market for medicinal and aromatic plants in India in order to assist the medicinal
plant industry to make informed decisions. (2) The biodiversity loss due to wild
harvesting of plants, and as an alternative the cultivation strategy of medicinal plants.
R. Kumari (*) • M.N.V. Prasad

Department of Plant Sciences, University of Hyderabad, 500046 Hyderabad, India
e-mail: rimabotany@gmail.com
226 R. Kumari and M.N.V. Prasad
(3) Main medicinal plant species having rich antioxidant potential. (4) Main
secondary metabolites of plant origin such as phenylpropanoids, terpenes, alkaloids,
and volatile oil, and other important metabolites containing high antioxidant level
used in human diet and health. (5) UV-B factors that enhance the quality of medicinal
plant by increasing the content of secondary bioactive products. (6) Secondary
metabolic pathways involving regulation of key enzymes, chalcone synthase, and
phenylalanine ammonia lyase. Understanding of UV-B responses on secondary
plant metabolites expand new opportunities for plant enriched in medicinal active
compounds.
Keywords Medicinal plants • UV-B • Antioxidants • Secondary metabolites

• Phenylpropanoids • Terpenoids • Alkaloids • Essential oil • Health benefits
List of Abbreviations
UV-B Ultraviolet-B
ROS Reactive oxygen species
NOx Nitric oxide
HIV Human immunodeficiency virus
SARS Severe acute respiratory syndrome
TIA Terpenoid indole alkaloids
PAL Phenylalanine ammonia lyase
C4H Cinnamate 4-hydroxylase
CHS Chalcone synthase
DFR Dihydroflavonol 4-reductase
DXS 1-deoxy-d-xylulose5-phosphate synthase
IPPI Isopentenyl diphosphate isomerase
GPP Geranyl diphosphate
FPPS Farnesyl diphosphate synthase
DAHP 3-deoxy-D-arabino-heptulosonate-7-phosphate
DPPH Diphenylpicrylhydrazyl
1 Introduction
During the past decade, therapeutic potential of medicinal and aromatic plants with
rich contents of secondary bioactive metabolites have received increased attention
for scientific investigation. Despite the progress in synthetic drug chemistry, plants still
constitute an important source of new products of medicinal value for pharmaceuticals,
used commercially either as an antioxidants or nutritional supplements (Zhang and
Björn 2009). Among the many medicinal plants routinely used as Ayurvedic medi-
cines in India, some of these plants have been featured in Fig. 1. Many of the herbal
Medicinal Plant Active Compounds Produced by UV-B Exposure 227
Fig. 1 Some selected Ayurvedic medicinal plants of India

drugs are simple synthetic modifications of these naturally plant derived secondary
metabolites. In recent years, increasing attention has been paid by consumers and
researchers to the health and nutritional aspects of metabolites such as phenolics,
vitamins, minerals and others. Secondary metabolites biosynthesis is a new tool in the
generation of novel and expensive natural products. Many of these phytocompounds;
phenols, flavonoids, steroids, alkaloids, terpenoids, anthocyanins, glycoscynates,
b-carotene, ascorbic acid, polyamines, synaptic ester, volatile oil, having high
antioxidant potential, be helpful in treatments of several oxidative stress related
disorders (Krishnaiah et al. 2007; Kaur and Arora 2009). These antioxidants mitigate
the risk of number of chronic and cardiovascular problems and other free radical
mediated diseases including neuro degeneration (Nambiar et al. 2010). Finding new
natural sources of antioxidant compounds with potential antiradical activity can be
useful to future therapy against various stress related health problems.
Increasing awareness of role of antioxidants and various active phyto- constituents
in medicinal plants is raising greater impetus for scientists to evaluate and determine
how the contents and quality of these natural products and their anti-oxidative poten-
tial can be maintained or even improved through different agro -technical processing
. In addition, Environmental factor also influence accumulation of key compounds.
Ultraviolet-B radiation (280–310 nm) is one factor, expected to modify the chemical
composition of plants and induces changes in level of secondary metabolites. Recently,
the knowledge about the treatment response of UV-B exposure on different plant
metabolites in terms of “health beneficial aspects” is exponentially increasing and has
led to novel insights, especially as far as medical, pharmacological and nutritive
aspects are concerned (Jansen et al. 2008). Evidently, based on findings by various
researchers this novel concept may be presented that, UV-B supplementation might be
beneficial for sustainable agriculture, food and nutritional industries for determining
the high commercial utilization of these natural medicinal products (Kumari et al.
2009a, c; Kumari and Agrawal 2010; Sun et al. 2010; Eichholz et al. 2011).
Rather then some researches goes on, there is limited understanding of the effect
on secondary plant metabolites such as phenolics and volatiles in different medicinal
plants, at all. The role and importance of UV-B on induction of phytocompounds in
today’s used medicinal plants needed to be explored as it may represent a new practical
approach to promote the quality and quantitative yield of metabolic product.
2 Global Trade, Demand and Supply of Medicinal

and Aromatic Plants from India
The world health organization has estimated that more than 80% of the world
population in developing countries depends primarily on herbal medicines for
basic healthcare needs (WHO 2002; Vines 2004). People rely more on natural
product derived from shrubs, herbs and trees of great medicinal value as herbal
medicine is safe and effective, at low cost with least and/ or no side effects. So on
in present scenario, medicinal and aromatic plants occupy a priming economic

position because of the continuous and increased demands of their products at
local, national and international markets. Further, there has been ever increasing
demand especially from developing country for more and more drug from plant
resources.
India is one of the major exporters of crude drugs mainly to six developed
countries; USA, Germany, France, Switzerland, U.K. and Japan, who share about
major export of total market. According to World Bank study, Global market for the
Botanical medicines is estimated as around US $60, and is projected to reach US$5
trillion by 2050 (WHO 2003). About 75–80% of the total exports of crude drugs
come from India (Shetty 2011). An overview of the global markets for medicinal
and aromatic plants from India, supply and international demand of these products
may expand information regarding opportunities of the markets for medicinal and
aromatic plants in India. This is to assist to make informed decisions on the develop-
ment of their medicinal plant industry.
3 Commercial Cultivation of Medicinal and Aromatic Plants
The growing demand of plant-based medicines, health products, pharmaceuticals,

food supplements, in etc., is putting an unsustainable threatening on forests resulted
in increased hazard and extinction of several species, because of overexploitation of
wild plant populations. Forests contribute to more than 90% of the medicinal plants
used for manufacturing medicines. Large-scale wild harvesting from the forests has
caused serious depletion of a number of economically-important species, loss of
genetic biodiversity, environmental degradation. According to estimate, more than
4,000 species of medicinal plants are globally threatened, largely due to commercial
over-harvesting to meet the demand from pharmaceutical industries (Schippmann
et al. 2002). So on, conservation of medicinal plants is needed for a number of
reasons. Firstly, they are an important source of natural ingredients used by the
manufacturers of modern pharmaceuticals. Second, its tremendous use as botanical
dietary supplements in past few decades, leads to continuous erosion of forest and
the forest product (Kala et al. 2006). Shifting from wild harvesting to cultivation
techniques appeared to be an important alternative strategy to overcome the problem
(Uniyal et al. 2000). In cultivation, agro-technological processing can be manipu-
lated in order to achieve desired products, increased yield or stable concentrations
of biologically active phytocompounds and reduced toxin level of botanical
products (Canter et al. 2005). Cultivation of medicinal and aromatic plants gives
scope to optimize yield and achieve a uniform, high quality product at cost of least
destruction (Wiersum et al. 2006). As a concluding remark, in this section, we
summarized the aspects of biodiversity loss related to wild harvesting of plants and
cultivation strategy of medicinal plants from the wild, to be – of better option, to
conserve valuable wild medicinal plants.
4 Therapeutic Use of Medicinal Plants as Antioxidants
The use of herbal drugs is widespread and serves as leading research for the
development of novel pharmacological agents. A large number of medicinal
plant species have already been tested for their potential biological, therapeutic
and pharmaceutical activities worldwide (Majhenic et al. 2007; Mata et al. 2007;
Wannissorn et al. 2005). Several studies from different countries have confirmed
their antioxidants activity in various plants having rich medicinal value
(Desmarchelier et al. 1998; Liu and Ng 2000; Souri et al. 2008; Ayoola et al.
2008). Fenglin et al. (2004) have screened the free radical scavenging activity of
Chinese medical woody plants showed that most of them are employed for their
effects on hemostasis, as anti-inflammatory, antimicrobial or for treatment of
dysentery and there medicinal uses may be directly linked to the content in tannins
and flavonoids and consequently to their free radical scavenging activities.
Zakaria (2007) identified the biochemical compounds such as Flavonoids, triter-
penes, saponins and tannins responsible for the observed antioxidant activity in
some herbal species; e.g. Muntingia calabura; Bauhinia purpurea; Dicranopteris
linearis; Melastoma malabathricum; Corchorus capsularis. Pourmorad et al.
(2006) noticed that greater amount of phenolic compounds relates with higher
radical scavenging effects in Mellilotus officinalis. In India also, several investi-
gators has been worked at this specific context in assessing the antioxidant capac-
ity of herbal medicine (Chanda and Dave 2009; Kaur et al. 2008; Ali et al. 2008).
Prakash et al. (2007) have identified the total phenol, antioxidant and free radical
scavenging activities of some Indian medicinal plants, e.g. Azadirachta indica,
Cassia fistula, Casuarina equisetifolia, Indigofera tinctoria, Lawsonia inermis
and Trewia nudiflora. These plants are having high phenolic content and antioxi-
dant activity. Kshirsagar and Upadhyay (2009) have studied the antioxidative
effects of 32 plant species from Tripura, northeastern India and observed that 16
of them e.g. Mitragyna rotundifolia, Schima walichii, Syzigium cerasoides and
others showed higher antioxidants potential depends on their dose dependent
activity. Jain et al. (2010) have studied on Alangium salvifolium and reported that
the extract from its roots have interesting potential free radical scavenging activity
for treatment of diseases. Patel et al. (2010) worked on antioxidant activity of
some selected medicinal plants from western region of India and screened the
high radical scavenging activity in the stem of Kigelia followed by leaf of
Hibiscus, Gemelia and Kigelia. Since the phyto-constituents and volatiles of
medicinal herbs have created renewed demand in their use by the public, explo-
rations of health benefits and antioxidant potential of these metabolites in the
prevention of oxidative stress problems is needed. This part described the summa-
rized view on antioxidant activity of various medicinal plants. Most of the
medicinal plants are rich antioxidants due to its different active ingredients that
determine its antioxidant potential.
5 Radical-Scavenging and Antioxidant Potential

of Secondary Metabolites
Oxidation process is one of the most important routes for producing free radicals
and reactive oxygen species (ROS), in food, drugs and living systems. These reac-
tive species exert can cause substantial biological damage; lipids, proteins and DNA
damage, leading to many chronic disorders in humans such as cancer, diabetes,
aging, and other degenerative diseases if excess ROS are not eliminated by antioxi-
dant defense system (Halliwell 1994; Petersen et al. 2005). Antioxidants plays
major role in reducing such free radical induced tissue injury and in overcoming
stress constraints, capable of acting as peroxide decomposers, singlet and triplet
oxygen quenchers, enzyme inhibitors and synergists (Edreva et al. 2008). Several
phytochemicals; phenolic compounds e.g. coumarins and quinones, nitrogen com-
pounds e.g. alkaloids, terpenoids, carotenoids e.g. beta-carotene, curcumin, ascor-
bic acid and flavonoids; orientin and vicinin have demonstrated to represents
significant antioxidant activity (Rice-Evans et al. 1995; Larson 1988).
5.1 Phenolic Compounds
Last few years, much interest has been attracted to use of natural and synthetic phyto-
phenolics as antioxidants, UV screens, anticancer, antivirus, anti-inflammatory,
wound healing, and antibacterial agents. Phenolics, belong to the phenylpropanoids
are major group of phytocompounds, having more then 10,000 groups of compounds.
They found to be derived mainly from phenylalanine and tyrosine. “Phenolics act as
multiple defense response as in photo-oxidative pigmentation such as anthocyanins,
flavonoids), antioxidation” (Alothman et al. 2009). The combination of antioxidant
abilities and scavenging reducing radicals makes the phytophenols ideal substrate for
balancing the cell redox status under oxidative stressed condition (Dai and Mumper
2010). Studies showed that the phytophenols can also quickly repair DNA radical
anions by an electron transfer reaction (Sperandio et al. 2002).
Among polyphenolic UV-B screening pigments, flavonoids aroused consider-
able interest because of their long had been shown beneficial effects on human
health. They have been reported to have antiviral, anti-allergic, anti-stress and
estrogenic activity, mitigating the risk of coronary heart diseases, amazing cardio-
protective effects through inhibition of peroxidation of low-density lipoproteins,
cholesterol, reduce platelet aggregation, or reduce ischemic damage (Nambiar
et al. 2010; Kim et al. 2008). Originally their good effects were thought to be due to
their “antioxidative” effect and chelating capacity (Atmani et al. 2009). Some research
evidences noticed the anti-tumor promoting activity of flavonoids via mechanisms as
kinase inhibition and apoptosis (cell cycle arrested) (Kanadaswami et al. 2005).
Anthocyanins are class of flavonoid compounds, which are widely distributed

plant polyphenols. Apart from their physiological roles in plants, anthocyanins
regarded as important components in human nutrition (Ross and Kasum 2002) that
supported by numerous studies, relating high positive correlation of fruit/vegetable
anthocyanin pigment content and increased antioxidant capacities (Stintzing and
Carle 2004). Anthocyanin functions in counteracting the negative effect of nitrogen
and oxygen reactive species (NOx), maintaining the redox homeostasis of biologi-
cal metabolism. All the effects of the anthocyanins listed above can be explained by
the antioxidant mechanisms, including hydrogen donation, metal chelation, and
protein binding (Kong et al. 2003).
Chlorogenic acid is one of the most important polyphenols in human diet. Like
other dietary polyphenols, chlorogenic acid is an antioxidant, produced by large
variety of fruits and vegetables, coffee plants in response to environmental stress
conditions. Some of the closely related phenolics such as ellagic acid, caffeic acid,
and ferulic acid, collectively called as Chlorogenic acid. Potentially beneficial
properties to humans are such as antioxidant, hypoglycaemic, antiviral and hepato-
protective (Farah and Donangelo 2006). These are also shown to have chemopre-
ventative activity in a number of animal tumor models (Gonthier et al. 2003).
5.2 Carotenoids
Carotenoids are naturally occurring pigmented compounds produced in many plants

and microorganisms. Their main biological function is the protection against oxida-
tive damage in plant defense system. Due to the unsaturated nature of the carote-
noids they are subject to changes mainly due to oxidation. The antioxidant properties
of carotenoids have been suggested as being the main mechanism by which they
afford their beneficial effects on human health. Carotenoids could possibly play an
important role as anticarcinogenic drug and in preventing several stress related
human disorder such as atherosclerosis, cardiovascular, and other degenerative
pathologies such as diabetes, Parkinson’s and Alzheimer’s diseases (González-
Gallego et al. 2007). The carotenoid ß-carotene is the primary source of vitamin A
in the human diet. Carotenoids may mediate their effects via other mechanisms such
as gap junction communication, cell growth regulation, modulating gene expres-
sion, immune response and as modulators of carcinogen metabolic pathway via
Phase I and II drug metabolizing enzymes (Paiva and Russell 1999).
5.3 Alkaloids
Alkaloids are a diverse group of low-molecular-weight, nitrogen-containing

compounds found in about 20% of plant species. Some alkaloids are poisonous,
while most are having medicinal value. Furthermore, alkaloids have an important
function in the immune systems of living organisms (Tadeusz 2007). The most
medicinally important alkaloids of plant origin includes vinblastine and taxol as
an anticancerous product, morphine an analgesics, colchicines; cell division sup-
pressant, and scopolamine as a sedative. Alkaloids interact with DNA or gene coding
enzymes, can influence electron chains in metabolism and modulate enzyme
activity. Cui et al. (2006) reported the antimicrobial, antimalarial, cytotoxic, and
anti human immunodeficiency (HIV) virus activities of the isoquinoline alkaloids
and explained that possible chemopreventive antitumor promoters are probably
related to their radical scavenging activity against diphenylpicrylhydrazyl (DPPH)
radical. b-carboline alkaloids showed strong activity against H2O2-induced oxidative
DNA damage, and their hydroxyl radical-scavenging property appears to contribute
to their antimutagenic and antigenotoxic effects, observed in yeast and mammalian
cells, respectively (Moura et al. 2007).
5.4 Terpenoids
Terpenoids are defined as secondary metabolites with molecular structures contain-

ing carbon backbones of isoprene (2-methylbuta- 1, 3-diene) units. These largest
classes of plant metabolites include many aromatic substances and essential oils
(Nassar et al. 2010). More than 40,000 terpenoids compounds have been identified
till with new compounds being discovered every year. Plants terpenoids are used
extensively for their aromatic quality. They play major role in traditional herbal
remedy, used as anti-bacterial, anti-neoplastic, anti-inflammatory, inhibition of cho-
lesterol synthesis and other pharmacological properties (Paduch et al. 2007;
Yamunadevi et al. 2011). Among most of the terpenoids produced by plants, few of
these are investigated in depth for pharmaceutical application for example artemisi-
nin and taxol for malaria and cancer treatment (Rai et al. 2011; Yamunadevi et al.
2011). In recent publication, Thoppil and Bishayee (2011) reviewed on the potential
role of naturally occurring different terpenoids in the chemoprevention and treat-
ment of liver tumors.
5.5 Phytosterols
Plant steroids, also called phytosterols, are isoprenoid-derived lipids and are inte-
gral components of the plant cell. Moreover, phytosterols have important pharma-
cological activities, including antitumor effects against lung, stomach, ovary and
breast cancer (Woyengo et al. 2009). Phytosterols limit absorption of cholesterol
from fat matrices into the human intestinal tract, decreases the incidence of cardio-
vascular diseases (Sanclemente et al. 2009). Phytosterols exerts anti-atherosclerotic,
anti-inflammatory and anti-oxidative activities.
5.6 Polyamines
Polyamines; putrescine, spermidine and spermine are a group of phytohormone-like

aliphatic amine natural compounds with aliphatic nitrogen structure. Polyamines
are small ubiquitous polycations, can modulates the function of important bio-
molecules under stress. These are modulator of stress regulated gene expression
and exhibit antioxidants property (Kuznetsov and Shevyakova 2007).
5.7 Glucosinolates
Glucosinolates are defense-related nitrogen- and sulphur-containing plant second-

ary metabolites, present mainly in species of Cruciferae family. Increase in these
aliphatic and indolyl glucosinolates, potentially have important benefits for human
health due to their anti-inflammatory and anti-tumorigenic properties (Gomes
et al. 2008; Kuhlmann and Muller 2009).
5.8 Essential Oil
Essential oils of aromatic plants have attracted a great deal of interest due to their
potential as a source of natural antioxidants and biologically active compounds
(Bozin et al. 2006; Tepe et al. 2007; Wannissorn et al. 2005). These active oil
gained attention of researchers for its multiple uses in phyto-therapeutics such anti-
inflammatory, antitumor, anti-hyperglycemic, anticarcinogenic, and as antioxidants,
(Eichholz et al. 2011). For treating intestinal and gastric problems, peppermint oil,
coriander oil, basil oil are used, lavender, eucalyptus and chamomile oils for insomnia
patients, muscles pain and aches (Wei and Shibamoto 2007), lemon grass oil for
treating relief in stress problems, calamus oil for improving intelligence power in
children (Kumari et al. 2009a, c).
In brief, this section of review, described the different major group of secondary
metabolites of plant origin such as phenylpropanoids, terpenes, alkaloids, volatile
oil as well as other important metabolites, containing high antioxidants level that
being used in human diet as nutrition supplement as well as important ingredients
of several medicinal products related to human health problem.
6 Needs of UV-B Research on Medicinal Compounds
The continuous increasing demand of the global market for aromatic and medicinal
plants led to the development of experimental cultivations where environmental
factors are of much concern which can improve the quality of products of the studied
plants in relation to their commercial value. In this prospective, UV-B researches on

medicinal plants have possessed a great deal of attention for researchers and phar-
maceutics, as they show the positive involvement of UV-B on quality and yield of
desired herbal products by inducing volatiles production and compounds of interest
of secondary metabolism; phenolics, flavonoids. It may be due to their special
consequence on biosynthetic processes as they play a variety of ecological roles.
They are having potency to enhance the level of terpenoids, phenylpropanoids, and
natural antioxidants (Kumari and Agrawal 2010).
7 Consequences of UV-B Stress: Phytocompound Induction
Application of abiotic stress such as moderate UV-B treatment can induce distinct
changes in the plant’s secondary metabolism. UV-B induction of different second-
ary metabolites is represented in Fig. 2. Ultraviolet supplementation has been
proposed to increase levels in tocopherol (Higashio et al. 2007), ascorbic acid and
flavonoids in some medicinal and aromatic plants (Kumari and Agrawal 2010).
Hagen et al. (2007) have evaluated the effects of postharvest UV-B irradiation with
visible light and reported the increased value on phenolic contents and evaluated
Shikimates Aromatic amino

acids
Terpenoids
(Salycilates,
(Terpenes gluconsinates
Steroids; (Phenylpropanoids) Glucocynolates,
Carotenoids) Flavonoids Indoles)
anthocyanin,
Lignins, tannins)
Pyruvate
Plant metabolism
Nutritional value/ human health

Phytoactive metabolites
+flavonoids,
induction + carotenoids,
+Amino acids,
+Terpenoids,
+ +Glucocynolates,
Defense signal
+Polyamines,
+Vitamines,
UV-B +Steroids, Ascorbic acid
stress + various nutrients

tocopherol
other medicinal products
Fig. 2 UV-B induction of different metabolites in plants

content of anthocyanins, quercetin glycosides, chlorogenic acid and ascorbic acid

including oxygen radical antioxidant capacity assay in apple fruit upon UV-B irra-
diation. UV-B treatment affects the important bioactive products in various medici-
nal plants as documented in Table 1.
UV-B study on secondary metabolites production is mostly centered on UV-B
screening “phenolics” compounds. These phytochemicals have been positively
linked to several health-promoting properties such as anticancerogenic and antioxi-
dative characteristics (Holst and Williamson 2004; Schreiner et al. 2009). Phenolics
mitigate the effects of oxidative stress by mediating the redox status (Kumari et al.
2009c). Studies reported an increase of phenolic compounds and corresponding
antioxidant activity, as in sweet flag (Kumari et al. 2009c), blueberries (Eichholz
et al. 2011) etc. Phenolics can act as UV-protectors e.g. because of their absorption
of light between 270 and 290 nm. UV-B research on rosamarinic acid, a bioactive
compounds of medicinal plant Rosmarinus officinalis (Rosemary plants) showed
higher yield of rosamarinic acid, when plants subjected to UV-B exposure (Luis
et al. 2007). Rosmarinic acid a naturally-occurring and potent phenolic compound
antioxidant plays a role in modulating inflammatory diseases including allergies,
asthma and atherosclerosis and anti-inflammatory properties.
Flavonoids are the major one UV-B-regulated phenylpropanoid derivatives, rap-
idly induced by UV-B exposure. Flavonoids stabilize and protect the lipid phase of
the thylakoid membrane, and are quenchers of the excited triplet state of chlorophyll
and singlet oxygen generated under excessive stress (Agrawal and Rathore 2007).
UV-B response on it’s biosynthesis and regulation have been very thoroughly
explored. Two groups of genes are required for flavonoids biosynthesis: structural
genes that encode the enzymes that directly participate in flavonoid biosynthesis,
and transcription factors that regulate the expression of the structural genes and the
accumulation of flavonoids. UV-B radiation stimulates the response of regulatory
key enzyme “Phenylalanine ammonia lyase” that’s an important enzyme of flavonoid
biosynthesis pathway and transcriptionally induced by UV-treatment (Ravindran
et al. 2010).
Some researches noticed the response of UV-B on glucosinolate metabolism,
as genes related to the biosynthesis of flavonoids, glucosinolates and terpenoids
metabolism were differently expressed after UV-B radiation (Hectors et al. 2007;
Wang et al. 2011). Flavonoid and glucosinolate induction depend partly on the
same signaling pathways, which involve jasmonic acid (Mackerness and Thomas
1999; Textor and Gershenzon 2009). UV-B application increased the aromatic
glucosinolate glucotropaeolin concentration up to sixfold in comparison to the
control plants in case study on short-term and moderate UV-B radiation effects on
a herbal medicinal plant nasturtium (Tropaeolum majus) (Schreiner et al. 2009).
Wang et al. (2011) investigated the contents of glucosinolates and the expression
of related genes in response to enhanced UV-B radiation and reported that when
Arabidopsis leaves were irradiated by acute UV-B (1.55 W·m−2) for a short time
(1 h), it induced the production of glucosinolates, however over exposure (12 h)
inhibited the expression of glucosinolate biosynthetic genes and glucosinolate
contents decreased.
Table 1 Effect of Ultraviolet-B (UV-B) radiation on secondary metabolites in some plants showing exampled compounds and its medicinal values
Plant name Medicinal property Active ingredients UV-B dose UV-B induced References
−2 −1
Acorus calamus L. antispasmodic, carmina- ß-asarone linalool +1.8 kJ m d Decrease in b-asarone Kumari et al.
tive, anthelminthic, aristolene, aristolene percentage (2009c)
Family: Acoraceae Hypotensive, caryophyllene No change in linalool
depressant carvacrol, phenolics Increase in aristolene,
p-cymene caryophyllene
oxide, carvacrol
Increased after UV-B
exposure
Artemisia annua L. antimalarial, anti-tumor, artemisinin +4.2 kJ m−2 d−1 Increased by 10.5% under Rai et al. (2011)
Family: Asteraceae anti-cancerous UV-B treatment
Camellia sinensis L. cardioprotective, Tea catechins Elevated epigallocatechin Zheng et al.
Family: Theaceae chemoprotective, Anti gallate level in tea leaf by (2008)
allergic artificial supplementation
of UV-B
Catharanthus roseus Anti cancerous, Strictosidine, Serpentine, 28 × 10−5 kJ/cm2 Increased respectively by Ouwerkerk et al.
Family: Apocynaceae Anti-diabetic Tabersonine, 161%, (1999)
Catharanthine, 8–14%
Vindoline, Vinblastine 21–79%
8–14%
8–14%
8–14%
***
Catharanthus roseus L. Catharanthine, vindoline 1.26 mW/cm2 Threefold and 12-fold Ramani and
Family: Apocynaceae increase respectively, on Jayabaskaran
treatment with 5-min (2008)
UV-B irradiation
***
Catharanthus roseus L. lochnericine serpentine, – Significant increases after Binder et al.
Family: Apocynaceae ajmalicine 20 min UV-B exposure (2009)
rhammericine Decrease in rhammericine
(continued)
Table 1 (continued)
−2 −1
Cymbopogon citratus Anti-depressant, Z-citral (terpenoids) +1.8 kJ m d Up to twofold increase at Kumari et al.
(D.C.) Stapf Anti-inflammatory, UV-B exposure (2009a, b, c)
Family: Poaceae Antipyretic
Genista Tinctoria L. hypoglycemic, Isoflavonols: genistin, – isoflavonols formed at higher Tumova and
Family: Fabaceae antiinflammatory genistein, daidzein, levels after UV irradiation Tuma (2011)
antiulcer, spasmolytic Biochanin-A
antioxidant,
estrogenic
Ginkgo biloba L. antimicrobial, neuropro- flavonoids Specific: 82.9 mWs·cm−2 Increased up to 57.2%trends Sun et al. (2010)
Family: Ginkgoaceae tector, anti-mutational, quercetin, kaempferol of UV-B induction in
and anticancer activity isorhamnetin quercetin, kaempferol,
glycosides and isorhamnetin
glycosides
Glycyrrhiza Uralensis L. Anti-tumor property Glycyrrhizin 1.13 W/cm2 UV-B stress stimulates Afreen et al.
Family: Fabaceae glycyrrhizin (2005)
concentration
Hypericum antidepressive properties Flavonoids, tannins and sUV-B Increase in level of Germ et al.
perforatum L. hypericin flavonoids, tannins at (2010)
Family: Clusiaceae supplemental UV-B dose
***
Mentha spicata L. Chem: I Piperitone sUV-B simulating 15% No effect of UV-B on Karousou et al.
I & II chemotype Chem: II; limonene, ozone depletion qualitative composition of (1998)
Family: Lamiaceae Carvone, component in both
dehydrocarvone chemotype
Mentha piperita L. Carminative, Menthone menthol 0.6 W m−2 Improve oil quality in terms Behn et al.
decongestant of enhanced rate of (2010)
Family: Lamiaceae Expectorant, relaxing menthone to menthol
effect on skin conversion
Ocimum basilicum Antioxidant, anticancer, Linalool, b ocimene UV-B enhancing effect on Johnson et al.
Family: Lamiaceae anti microbe methyleugenol, these phenylpropanoids (1999)
Eugenol, and terpenoids about
fourfold increase in
eugenol
***
Ocimum basilicum L. 1,8-cineole germacrene C, PAR, UV-B 41% and 71.4% increment Nitz and
Family: Lamiaceae linalool eugenol (2.0) kJ m−2 d−1 approx. 3 times at UV-B Schnitzler
(2004)
Ocimum basilicum L. Antioxidant, anticancer, 1, 8-cineole, Linalool, 222.6 W/m2 Significant increase in level Chang et al.
Family: Lamiaceae anti microbe eugenol. of these constituents at (2009)
UV-B exposure
Perilla frutescens L. anti-inflammatory, perillaldehyde limonene 0, 0.21, 0.88 W m−2 UV-B suppressed anthocya- Nishimura et al.
Family: Lamiaceae anti-pyretic nin production, No effect (2008)
on active components
Rosmarinus officinalis L. Antioxidant, anti-bacte- Rosmarinic and carnosic 5.4 and 31 kJ m−2 d−1 Both UV-B radiation Luis et al. (2007)
Family: Lamiaceae rial property acids treatments increased the
concentrations of caffeic,
carsonic, rosmarinic acid
Taxus chinensis L. antitumor agent Taxol 3.38 mw cm−2 nm−1 Increased taxol content at Zu et al. (2010)
Family: Taxaceae supplemental UV-B
treatment
Tropaeolum majus Anti-microbial effect Glucotropaeolin, Phenolic 0.075 and 0.15 W h m−2 UV-B application increased Schreiner et al.
Family: Tropaeolaceae the glucotropaeolin (2009)
concentration up to sixfold
in compare to control
Increased at UV-B exposure
Vaccinium corymbosum Antioxidant, Anti- micro- Total phenolics low (0.075), high UV-B exposure resulted in an Eichholz et al.
Family: Ericaceae bial, anticarcinogen C6-aldehydes: (0.15), UV-B dose, increase in phenolic (2011)
terpenes, ketones Wm−2 compounds, Increased
aldehydes
***
Represent as descript above; C represents: control, UV-B ultraviolet-B, PAR photosynthetic active radiation
Scientific interest in UV-B research on anthocyanin pigments have been well

documented because of their possible health benefits as dietary antioxidants.
Anthocyanins, a class of flavonoids is believed to increase the antioxidant potential
of plants in order to uphold the balanced physiological status in tissues under UV-B
stressed factor. The elevated levels of the transcripts of Pal, Dfr and Ans were
accompanied by an increase in the anthocyanin pigments (Guo et al. 2008). In several
medicinal/fruited cultivars, anthocyanin accumulation is induced by light in the
UV-B region (wavelengths from 280 to 320 nm) when applied in combination with
solar infrared radiation (Arakawa 1988). Ubi et al. (2006) reported the increased
accumulation of anthocyanin pigments in apple fruit skin by inducing the expression
of the anthocyanin biosynthetic genes, especially chalcone synthase, anthocyanidin
synthase and anthocyanin −3-O-glucosyl transferase genes.
“Coumarins and furocoumarins have been of particular interest because they are
induced by UV radiation”, Psoralens is a naturally occurring furocoumarins) absorb
strongly within the ultraviolet range (200–320 nm) band. Their photosensitizing prop-
erty makes them clinically effective in treatment of various dermatological ailments in
human being, as vitilago, leucoderma, leprosy and psoriasis, inducing re-pigmentation
(melanocytes formation). Psoralens are reported to be influenced by changes in the
UV-B environment in Abelmoschus esculentus (Kumari et al. 2009b). Coumarins and
furanocoumarins also absorb strongly in the UV wave bands and increase with
enhanced UV-B radiation in Pastinaca sativa (Zangerl and Berenbaum 1987).
UV-B promotion of terpenoids observes particularly in members of the Lamiaceae
family (Johnson et al. 1999; Karousou et al. 1998; Maffei and Scannerini 2000; Behn
et al. 2010). A study on Rosemary plant, reported UV-B radiation effect on diter-
penes carnosic acid and carnosol (Luis et al. 2007). Johnson et al. (1999) reported the
accumulation of terpenoid compounds at seedlings stage (five leaf growth) in basil
leaves. Glycyrrhizic acid, a triterpenoid saponin, were found to be increased at opti-
mum UV treatment while with exposure to high UV doses resulting in decreased
glycyrrhizin accumulation (Afreen et al. 2005). Glycyrrhizin is major bioactive com-
ponents of Glycyrrhiza uralensis (liquorice) having anti-tumor effect and inhibition
of carcinogenicity by the activation of hepatotoxic metabolites (Chan et al. 2003),
high activity in inhibiting replication of human immunodeficiency virus (HIV-1) and
severe acute respiratory syndrome (SARS) -associated virus (Afreen et al. 2005), and
exhibits a number of other pharmacological effects (Zhang and Ye 2009).
In Catharanthus roseus, terpenoid indole alkaloids produced in as part of its
secondary metabolism. This is supported by the observations that UV light induces
the formation of dimeric terpenoid indole alkaloids in C. roseus leaves (Hirata et al.
1993; Ouwerkerk et al. 1999). Ramani and Chelliah (2007) reported the significant
roles of UV-B induced signaling pathway on transcription of terpenoid indole alka-
loids biosynthetic genes encoding tryptophan decarboxylase (Tdc) and strictosidine
synthase (Str) expression leading to increased production of catharanthine in
Catharanthus roseus cell cultures. An anti-inflammatory monoterpene- indole alka-
loid brachycerine in plants of Psychotria brachyceras (Rubiaceae) induced almost
doubled by UV-B radiation (Gregianini et al. 2003). In study on Taxus chinensis,
variety mairei, Zu et al. (2010) found the increased content of a alkaloid “taxol” in
plants exposed to increased UV-B radiation. Taxol could significantly decrease the
concentration of free radicals generated by the classical fenton reaction and exists
as an important anti-oxidant substance. UV induction of alkaloid biosynthesis may
be useful for pharmaceutical purpose.
Polyamines may play role in protecting plants against UV-B stress (Kramer
et al. 1991). Polyamine acts as stress messengers in plant responses to stress signals
(Gill and Tuteja 2010). Polyamines protect against radiation-induced oxidative
stress (Deutsch et al. 2005). It plays main role in regulating structure and function
of photosynthetic apparatus, in avoiding cellular damage of lipids, proteins and
DNA of plants under UV-B stress (Sung et al. 2008; Agrawal et al. 2009).
From the nutritional point of view, UV-B irradiation has even been beneficial
in terms of increasing the content of potentially health-promoting stilbene
“resveratrol”. Resveratrol (3,5,4¢-trihydroxy-trans-stilbene) is produced by certain
plants as a defense mechanism. The biological activity of resveratrol as an anticarci-
nogenic and an antioxidant has been reported in graps (Jang et al. 1997). A series of
involved enzymes, such as phenylalanine ammonia-lyase, cinnamate 4-hydroxylase,
p-coumaroyl-CoA ligase and stilbene synthase involved in the resveratrol synthesis
that shown to be up-regulated by UV-B irradiation (Li et al. 2008).
In recent years some studies have been carried out concerning the impact of
enhanced UV-B radiation on hypericin concentrations and reporting that concentra-
tions of this bioactive substance can be altered by UV-B (Germ et al. 2010).
Hypericin is a component of the inducible plant defense response of Hypericum
perforatum against fungal pathogens (Cirak et al. 2005).
Concluding remark of this part; UV-B factors enhance the quality of medicinal
plant via enhancing content of secondary bioactive products in turms of induction
of phenolics, flavonoids and other secondary products,
8 UV-B Induction of Essential Oil Yield in Medicinal

and Aromatic Plants
The commercial value of an aromatic and medicinal plant could be reflected by the
yield and composition of their essential oils. An essential oil is a concentrated, hydro-
phobic liquid containing volatile aroma compounds having characteristic fragrances
in some plants. In several recent and earlier reports on medicinal plants, UV-B radia-
tion is reported to be beneficial in improving the quality of various aromatic plants,
showing it’s positive role in volatile production as well as inducing changes in chem-
ical composition of its essential oil. An aggregate accent of documented works done
so far on the impact of UV-B radiation on volatile oil yield and its qualitative compo-
sition in some medicinal plants are shown in Table 2. The contribution of UV-B
radiation to stimulation of volatile production and changes in active constituents
composition has been examined experimentally by various other researchers as
reported in Ocimum basilicum (Johnson et al. 1999; Nitz and Schnitzler 2004; Chang
et al. 2009), Mentha spicata (Karousou et al. 1998), Mentha piperita (Maffei and
Scannerini 2000); Glycyrrhiza uralensis (Afreen et al. 2005). Dolzhenko et al. (2010)
reported in study on peppermint that UV-B radiation prompted the higher content of
Table 2 Ultraviolet-B (UV-B) induced response on volatile oil yield and qualitative and quantitative change in essential oil constituents in various medicinal plants
from different countries
Photosynthetic
active radiation
(mmol phot. m−2 Experimental
Medicinal plants UV-B (kJm−2 d−1) sec−1) condition Supplemental UV-B induced response Reference Country
Acorus calamus L. Control (ambient) 1,100–1,200 Field condition Increment in essential oil yield by 35% Kumari et al. India
sUV-B (+1.8) Significant variation in qualitative (2009c)
quantitative composition of oil
decrease in b- asarone (8.5%), linalool
and germacrene-D.
Increase in aristolene (47.6%), caryophyl-
lene oxide (66.7%), p-cymene and
carvacrol ( up to detectable limit) and
other component
Cymbopogon Control (ambient) 1,100–1,200 Field condition Essential oil yield increased by 25.7% by Kumari et al. India
citratus L. sUV-B (+1.8) treatment of sUV-B. (2009a)
Major comounds of C. citratus oil: Z-
citral; increased more than doubled
after treatments
Mentha spicata L. Control (ambient) Near Ambient Field condition UV-B stimulated oil production in chem. I. Karousou et al. Greece
sUV-B (15% No change in qualitative composition of (1998)
Chemotype I and ozone oil.
II depletion) Slight variation in quantitative composi-
tion in chem I.
Mentha piperita L. Control (−UV-B) – Green house Increase in essential oil production Maffei and Italy
UV-B Variation in qualitative composition no Scannerini
quantitative change in oil constituents (2000)
Mentha piperita L. Control (−UV-B) 550 and 1,150 Growth At high PAR level, Increase in essential oil Behn et al. Germany
UV-B chamber yield at UV-B (2010)
(0.6 W m-2) Improved oil quality in terms of an
enhanced menthone to menthol
conversion
Ocimum Control (−UV-B), Ambient day Green house At early stage, no change in volatile yield Johnson et al. China
basilicum L UV-B light In five leaf stage of plant: fourfold (1999)
enhancing effect of UV-B on total
volatile content
Increase in phenylpropanoids and
terpenoids
High increase in eugenol (5 times).
Ocimum Control (−UV-B), – Green house Increase in essential oil yield (approx >2 Ioannidis et al. China
basilicum L. UV-B normal times) non-significant variation in (2002)
dose qualitative or quantitative composition
of volatiles constituents.
No change in number of oil glands
Expansion and fulfilling of peltate oil sacs,
increased aroma due to bursting of oil
sacs after sUV-B exposure
Ocimum Control (−UV-B), – Green house Increase in essential oil level: (16% in Nitz and Germany
basilicum L. PAR variant, PAR; more then 50% in sUV-B). Schnitzler
UV-B variant sUV-B induced changes in oil constituents: (2004)
Increase in 1,8-cineole (41%), germacrene
(71.4%), linalool and eugenol (approx.
3 times) decrease in methyleugenol
Increase in phenolics
Ocimum Control (ambient) Ambient day Field condition increased turgidity of oil glands increase in Kumari and India
basilicum L. UV-B (+1.8) light essential oil yield by 42% significant Agrawal
increase in b-caryophyllene, germac- (2010)
rene-D, ethyl linoleolate, b – elemene ,
camphenol
sUV-B supplemental UV-B above ambient, −UV-B UV-B excluded, – information not available
Fig. 3 UV-B stress response on plant metabolism; abiotic stress (e.g., from UV-B radiation) –
initiates signal cascades that trigger defense-related genes transcription. Resulting metabolic shift-
ing to secondary metabolism lead to generates higher secondary products as adaptive responses
menthol and phenolic compounds as the modulation of expression of some specific

genes involved in essential oil biogenesis (which show menthol production) was usu-
ally up-regulated by UV-B irradiation. Moderate and low dose UV-B supplementa-
tion are reported to increased the essential oil yield in a important Indian medicinal
plant Acorus calamus and shown to improve it’s health value by reducing the content
of potentially toxic constituents b- asarone (Kumari et al. 2009c). Another study on
Cymbopogon citratus also shown enhancement in it,s medicinal benefits by procuring
the higher content of z-citral (a major product) in essential oil as well as enhancing
it’s yield (Kumari et al. 2009a). Ioannidis et al. (2002) reported the requirement of
UV-B for normal development of oil glands in basil leaves.
Concluding remark; UV-B promotes volatile yield as well as induced changes in
it’s composition, that might proves to useful for enhancing the quality of medical
products.
9 UV-B Regulation of Secondary Metabolic Pathways
Plant secondary metabolites are important determinants of plant stress responses

(Jansen et al. 2008) (Fig. 3). UV-B radiation is linked to CO2 assimilation rate to
which the carbon skeleton is diverted from the primary metabolism to the secondary
one (Kasige and Takashi 2009). Therefore, UV-B regulation of carbohydrate

assimilation and secondary metabolism must be metabolic linked (Interdonato
et al. 2011). Most of the stress inducible enzymes such as glutathione-S transferase
and phenylalanine ammonium lyase play an important role in triggering secondary
metabolites synthesis. Phenylalanine ammonium lyase along with chalcone
synthase and other enzyme involve in allocating significant amount of carbon from
phenyl alanine into the biosynthesis of several important secondary metabolites
(Singh et al. 2009). Plant cytocromes P450s; cytochrome p450 family proteins and
p450 monooxygenase involved in catalyzing the metabolic reactions of primary and
secondary pathways to produce the biochemical products such as phenylpropanoids,
alkaloids, terpenoids, lipids, cyanogenic glycosides, and glucosinolates (Agrawal
et al. 2009). However, the exact molecular basis of UV-B signaling cascades leading
to increased production of secondary metabolites of plant cell is largely unknown.
These major routes for the production of secondary metabolites in plants are:
(1) shikimate pathway for phenylpropanoid compounds; flavonoids and anthocyanins
and isoprenoid route of (2) terpenoids and (3) alkaloids biosynthetic pathway. The pre-
sent literature describe with emphasis on the role of UV-B on these metabolic pathway.
9.1 Shikimate Pathway
Many of plant-derived phenylpropanoids such as flavonoids, isoflavonoids, antho-

cyanins, coumarines, and lignans are secondary products of shikimate metabolism
(Korkina 2007). Expression levels of several genes encoding key enzymes of
secondary biosynthetic pathway are being elevated in a coordinated trend at UV-B
and other environmental stress signals (Long and Jenkins 1998). Figure 4 showed
the diagrammatic representation of different metabolic steps involves in shikimate
pathway of phenylpropanoids biosynthesis and it’s regulation in response to UV-B
radiation. The first enzyme of the shikimate pathway “3-deoxy-D4 arabino-
heptulosonate-7-phosphate synthase” (DAHP synthase) that catalyzes the conden-
sation of phosphoenolpyruvate and erythrose-4-phosphate to yield 3-deoxy-
D-arabino-heptulosonate-7-phosphate (DAHP) is reportedly induced by UV-B
irradiation (Ramani et al. 2010). It proposed that selective transcription of phenyla-
lanine ammonia lyase, chalcone synthase, cinnamate 4-hydroxylase and 4-coumarate:
coenzyme A ligase genes, coding functional enzymatic variants, may control the
synthesis of phenylpropanoid products also being induced by UV-B stress factor.
9.2 Isoprenoids Pathway
The stimulation of terpene biosynthesis by oxidative stress is well known (Beaulieu

2007). The biosynthesis of terpenes starts with isopreneoid unit with precursors of
isopentenyl and dimethylallyl diphosphate. Activation of two independent pathway
Fig. 4 Schematic representation of UV-B regulation of phenylpropanoids pathway of

flavonoid biosynthesis. UV-B stress interferes with metabolic processes (particularly enzyme
activity). UVR8 controls multiple gene encoding the principle enzymes of flavonoid biosynthesis.
It acts primarily on phenylalanine ammonia lyase (PAL) and chalcone synthase (CHS); key
enzymes of flavonoids biosynthetic metabolism
cytosolic acetyl mevalonate, that provides precursor for sesquiterpene and sterols
and/or plastidial 1-desoxy-D-xylose-5-phosphate/methyl D-erythritol phosphate for
isopentenyl and dimethylallyl diphosphate precursors (Hampel et al. 2005). In a
recent publication on grapevine leaves, Gil et al. (2012) suggest that UV-B modu-
lates the metabolism of terpenes leading to specific responses according to fluence
rate of UV-B. Relatively low UV-B irradiation induces the de novo synthesis of
enzymes of terpene cytosolic mevalonic acid pathway leading to production of ter-
penes, triterpenes and sterols, while high UV-B irradiance promoted the production
of plastidic diterpenes via the methylerytritol phosphate pathway. The effect of
increased UV-B irradiation on the expression of genes involved in early steps of
terpenoid biosynthesis was reported in peppermint oil by Dolzhenko et al. (2010).
“They observed that UV-B irradiation increased the content of some monoterpenes
e.g. methol) and sesquiterpenes” e.g. b-caryophyllene and germacrene-D which
correlates with induced up-regulation of genes involved such as Dxs, Ippi, Gpps,
and Fpps. Alkaloid biosynthesis in plants is also tightly controlled in response to

UV-B stress. Tryptophan decarboxylase, a key enzyme in biosynthesis of the
precursor tryptamine needed for monoterpene indole alkaloid production, can
be strongly induced by UV-B light at the level of transcription (Ouwerkerk
et al. 1999).
At all, basic concept behind the activation of combitorial secondary metabolic
pathways is due to genetic up regulation of specific key enzymes; chalcone synthase,
phenylalanine ammonia lyase. Plant precursors from their own primary and
secondary metabolism are metabolized to the desired secondary product due to the
up-regulation/ expression of specific/ desired genes in response to UV-B irradiation.
10 Applicability of UV-B on Metabolites Enrichments:

Future Perspectives
The valuable pharmacological properties of phytocompounds limited with their low

level of production by natural sources. Active researches into novel strategy implied
for promoting metabolites production, will definitely be of great importance in
present and future in terms of their commercial value. Additional knowledge on
specific effect of light quality on metabolites induction will help us in designing
the greenhouse light environment to obtain plants with enhanced phytochemical
concentrations. The approaches discussed above –health value of various metabolites,
UV-B induction of these phytocompounds, secondary biosynthetic pathway, and the
UV-B induction of regulatory genes – are likely to be used in combination with new
high-throughput strategies to identify the most preferred route correlating with
desired metabolite biosynthesis in different plants under UV-B exposure experi-
ments. Provided that models are validated to ensure they reflect reality in targeted
increase in phytochemicals, this information will be of great value for identifying
and increasing the health promoting effects of medicinal plants and predicting
the cropping strategies on these plants in a changing global climate. This baseline
information generated will be useful for the successful implementation of UV
technology on a pilot scale, to benefit health conscious consumers, attributed to
concurrent increase in different health-related constituents.
Future research studies are warranted to evaluate the effects of different doses/
exposure time of UV-B on the contents and heath qualities of products in multiple
cases which have not yet been investigated. Such data sets will provide a wealth of
information to ecologists, plant breeders and agronomists.
10.1 Commercial Relevance
The fact, that solar UV-B regulates the secondary pathway of plants and improves
plant metabolic functioning especially in aspects of food quality, pharmaceutical
properties, pest and disease resistance. In this regard, UV-B supplementation
processing may be proven as a useful tool for plant manipulation in aspects of

optimizes the nutritional/health value of herbal drugs and subsequently generate
new opportunities by farmers and processors for achieving its rich quality and
increased essential oil yield.
The information on health sensory constituents and antioxidant potential of
medicinal and aromatic plants should be ever encouraged as it showed role on
reduction of the risk of diseases. Phytochemistry study, as reported on medicinal and
aromatic plants showed a positive induction of antioxidant compounds, such as
polyphenols and flavonoids, essential oil yield and qualitative composition of oil in
response to UV-B stressing. This enhancing effect of UV-B may be an interesting to
follow in studying the phytotherapeutic activity of medicinal plants. In this review
we provide an overview of medicinal property of various secondary metabolites, its
responses to UV-B irradiation stimuli as well as reported pathway of secondary
metabolic mechanism. Overall, the present article has added to our understanding
of statutes and prospective of medicinal and aromatic plants, its health aspects on
human life as well as positive response of UV-B to enhance nutritional value of
plants via inducing possible enhancement in various health promoting constituents.
11 Conclusion
Short-term and moderate UV-B radiation seems to be a safe alternative as targeted

treatment in medicinal and aromatic plants attributed to concurrent increase in
different health-related constituents. In terms of sensory attributes as well as health
benefits of valuable products of medicinal importance, these studies might contribute
to provide a convenience and feasible approach to enhance the yield and qualitative
value of these phytochempounds and subsequently generate new opportunities for
growers and processors, for achieving the health-oriented herbal product market.
In this respect, this may provide an innovation approach in pharmaceuticals drug
technology. Hence, to determine relevant metabolic interactions between specific
target metabolites and UV-B radiation may be a challenging task.
Acknowledgement The author Rima Kumari acknowledges Dr. D.S. Kothari Post Doctoral
fellowship from University Grant Commission for financial support. Thanks are also due to UGC,
New Delhi for providing research grant “Precision stressing by UV-B radiation to improve the
quality of Coriander and Trigonella, Ref. F.No. 41-389/2012
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Assessing the Environmental Benefits
of Compost Use-on-Land through an LCA
Perspective
Julia Martínez-Blanco, Cristina Lazcano, Alessio Boldrin,

Pere Muñoz, Joan Rieradevall, Jacob Møller, Assumpció Antón,
and Thomas H. Christensen
Abstract Due to increasing compost use in agriculture, there is an urgent need to

evaluate compost benefits and impacts versus other fertilizers. Here we review the
recent progress made in the quantification of positive effects associated with com-
post use on land using life cycle assessment (LCA), an internationally recognised
environmental tool. Nine environmental benefits were identified in an extensive lit-
erature review: nutrient supply, carbon sequestration, weed pest and disease sup-
pression, increase in crop yield, decreased soil erosion, retention of soil moisture,
increased soil workability, enhanced soil biological properties and biodiversity, and
gain in crop nutritional quality. Quantitative figures for each benefit were drawn
from the literature and classified into short-term: less than 1 year; mid-term: less
than 10 years and long-term: less than 100 years.
J. Martínez-Blanco (*)
Institute of Environmental Science and Technology (ICTA),
Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Spain
e-mail: julia.martinez@uab.cat
C. Lazcano
Centro Tecnológico del Mar – Fundación CETMAR, Vigo, Spain
A. Boldrin • J. Møller • T.H. Christensen
Department of Environmental Engineering, Technical University
of Denmark, Lyngby, Denmark
P. Muñoz
Institute of Agri-food Research and Technology (IRTA), Cabrils, Spain
J. Rieradevall
Institute of Environmental Science and Technology (ICTA),
Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Spain
Department of Chemical Engineering, Universitat
Autònoma de Barcelona, Cerdanyola del Vallès, Spain
A. Antón
Institute of Agri-food Research and Technology (IRTA), Cabrils, Spain
Departament d’Enginyeria Química, Universitat Rovira i Virgili (URV), Tarragona, Spain
256 J. Martínez-Blanco et al.
The review shows that 5–60% of the applied compost N is mineralized, depending
on the time frame. Mineralisation ranges from 35% to 100% for P and from 75% to
100% for K. Carbon sequestration rates are higher in the short term, up to 40% of
the applied C, decreasing over time to 16%. Impacts on crop yields vary from
decreases of 495% to increases of 52%. Compost increases 29–63% soil aggregate
stability, reducing 5–36% soil loss. Soil bulk density decreases of 0.7–20% after
compost application, potentially increasing soil workability. Also, water holding
capacity and plant available water can increase by 50% and 34% respectively. Data
on compost impacts on soil biodiversity is scarce and restricted to microorganisms.
Compost may decrease microbial diversity by 2% or increase it by 4%. Clear
impacts are also observed on microbial biomass, of 3.2–242% increase after com-
post application, and microbial activity: 43–344% increase. For weed, pest and dis-
ease suppression, along with nutritional content increase, quantitative figures could
not be provided, either because of lack of data or because the effects are very vari-
able and depend on specific local conditions. For soil erosion and soil moisture
content, effects could be quantitatively addressed, but suitable impact assessment
methodologies were not available. Additional impact categories dealing with
phosphorus resources, biodiversity, soil losses, and water depletion are needed for a
comprehensive assessment of compost application.
Keywords Sustainable agriculture • Environmental impact • Life cycle assessment

• Compost • Biowaste • Organic matter • C sequestration • Soil
List of Abbreviations
CLPP Community level physiological profiles

GHG Greenhouse gas
IPCC Intergovernmental panel on climate change
LCA Life cycle asessment
LCI Life cycle inventory
MSW Municipal solid waste
PCR-DGGE Denaturing gradient gel electrophoresis
SOC Soil organic carbon
SOM Soil organic matter
TRFLP Terminal restriction fragment length polymorphism
1 Introduction
There is increasing concern about soil interrelated environmental problems such as

soil degradation, desertification, erosion, and loss of fertility (European Commission
2006). These problems are partially consequence of the decline in organic matter
content in soils. A level of 2% of soil organic carbon (SOC) is commonly considered
Assessing the Environmental Benefits of Compost Use-on-Land… 257
Fig. 1 Municipal solid waste compost applied to a local field (Photo by Pere Muñoz)
desirable for maintaining good soil structure for agricultural activities (Van-Camp
et al. 2004). An estimated 45% of European soils have low (<2%) soil organic matter
(SOM) content, principally in southern Europe but also in areas of France, the UK,
and Germany (European Commission 2006).
A parallel concern is the massive generation of organic waste by human activities –
for instance, organic waste represents 30–40% of the municipal solid waste (MSW)
generated in the European Union (European Parliament 2010) –, which has led to the
proposal of several alternatives to avoid landfilling and promote recycling. Among
these alternatives, composting is one of the best-known processes. Composting
allows the stabilization and sanitation of organic waste through accelerated aerobic
decomposition under controlled conditions. In addition, depending on the quality of
the initial feedstock, the final compost can be used for agricultural purposes in a local
context (Fig. 1).
Several studies indicate that the use of compost on land may improve several
plant and soil parameters (Fig. 2), which would make compost an interesting fertilizing
option not only for agriculture but also for soil restoration purposes. Compost
addition increases SOM content, which enhances aggregation and stability, thereby
ameliorating soil structure (Diacono and Montemurro 2010). Stability of soil aggre-
gates prevents surface sealing, improves water infiltration, and enhances water hold-
ing capacity thus reducing runoff generation and soil erosion (ROU 2007). Moreover,
increasing SOM levels promotes carbon sequestration (Favoino and Hogg 2008;
Crop nutritional
Crop yield quality
Weed, pest and

disease suppression
Carbon
sequestration
Soil
workability
P Soil
erosion
N K
Soil moisture Nutrient Soil biological properties
content supply and biodiversity
Fig. 2 The nine benefits of compost application assessed in the paper
Marmo 2008). Other potential benefits of compost application are improved

biological activity (Bastida et al. 2008; Hargreaves et al. 2008), enhanced nutrient
availability (Boldrin et al. 2009), and the suppression of soil borne diseases
(Bonanomi et al. 2007). Furthermore, several authors have reported higher yields
with compost application and better quality of the harvested crops. Nevertheless,
the application of compost may also produce environmental and agronomic draw-
backs. The most relevant ones being gaseous and leachate emissions, immobiliza-
tion of soil mineral N, increase in salt content, and accumulation of heavy metals in
soil (Hargreaves et al. 2008). These issues are in general directly associated to the
quality of the feedstock waste used, the maturity of the compost, and the application
of the compost (e.g. crop rotation, soil type, etc.). Nevertheless in recent times, the
increasing societal concern about pollution, land degradation and resource con-
sumption is introducing other type of criteria such as environmental impacts and
benefits in the decision process. Decision tools are now forced by society and regu-
lations to be focused on environmental aspects.
Several tools are available to quantify positive and negative environmental
impacts of compost in a comprehensive manner. Among them, Life Cycle
Assessment (LCA) was promoted in different European directives as a robust
quantitative tool, and a keystone in decision making by producers and stakeholders.
LCA has been widely used for the assessment of the waste and agricultural sectors.
While the environmental impacts associated to compost production have been

successfully assessed in previous studies (Blengini 2008; Boldrin et al. 2010;
Hansen et al. 2006; Lundie and Peters 2005; Martínez-Blanco et al. 2010, 2011;
Meisterling et al. 2009; ROU 2007), assessment of most of the benefits of compost
on plant and soil have not been taken into account. Several recent studies address
the inclusion of compost benefits in a qualitative manner, recommending that
further research should be undertaken on the subject, (Boldrin et al. 2009; Favoino
and Hogg 2008; Hansen et al. 2006; Martínez-Blanco et al. 2011). Nevertheless,
carbon sequestration and nutrient supply are, to date, the only environmental
benefits taken into account when evaluating the environmental impacts of compost
application. Because of the modelling complexity, ROU (2007) is, to our knowl-
edge, the only study where an attempt was made to include most of the abovemen-
tioned effects within LCA of two Australian case studies. The results are however
only presented at the inventory stage and the obtained figures are not included
within the impact categories.
The main goal of this paper is to quantitatively address LCA modelling of
those positive effects traditionally associated to land application of compost origi-
nating from organic MSW and garden waste. This is done by (1) explaining some
of the main dynamics determining the considered environmental and agronomic
benefit (from now on benefit); (2) providing quantitative data (inventory) based on
a critical revision of state-of-the-art literature; (3) describing existing impact
methodologies and future challenges for assessing the benefits under a LCA per-
spective. Martínez-Blanco et al. (submitted 2012) overviews the same goals and
literature review.
2 Methodology
A comprehensive revision of the literature dealing with the effects of compost on

soil and plant growth was carried out between November 2010 and May 2011. The
literature revision was divided into two parts, following the goals of the paper: (1)
the potential benefits of compost application and (2) the current situation of the
inclusion of each of these benefits in LCA studies.
2.1 Literature Review on Potential Compost Benefits
The most relevant effects of compost on soil properties and plant growth were first
identified through a literature search. Subsequently, 90 articles (including both
reviews and case studies) were selected for further revision and collection of inven-
tory data. The following criteria were used:
• Data from organic MSW or biowaste (food and kitchen waste from households,
restaurants, caterers and retail premises, and comparable waste from food
processing plants) and green waste (coming from private gardens and public
parks) were taken into account.
• Field studies were selected, when possible, instead of pot and laboratory incubation
experiments.
• Compost used for agricultural purposes was in focus. However, land reclamation
or restoration studies were also taken into consideration when necessary.
• Articles published earlier than 1990 were not taken into account, when possible,
as they were replaced by better or newer studies.
Some of the studies included in the review did not fulfil one or more than one of
the previous criteria. These studies were still included as they constituted the only
scientific contribution in a specific area or because they were particularly relevant.
According to the literature revision, the effects were classified into short term
(1 year), mid term (1–10 years), and long term effects (10–100 years), depending on
the length of the studies. Thus, quantification of specific effects is associated with
the corresponding time frame.
Regarding the combination of compost, doses and experiment length, the major-
ity of the reviewed papers on the short-term considered a unique application,
while periodically applications are taken into account for longer studies. For the
homogenization of the data, the annually proportional rates were considered and
therefore in these cases the cumulative effects of compost application, rather than the
residual effect were reported. All the publications included for each of the assessed
compost benefits were summarized in tables. The more relevant variables – for
instance the type of composted biowaste and the length of the experiment – were
defined for each study, as well as the corresponding parameters used to measure the
benefit. According to the specificity of each compost benefit, either average values
or range values were stated when possible.
2.2 Life Cycle Asessment of Compost Use on Land
With regards to the LCA quantification and assessment methodologies, The ranges
obtained in the literature review for each of the compost benefits were used as start-
ing point for the LCA quantification and assessment methodologies. Subsequently
a broad revision of the current status of the corresponding assessment methodolo-
gies was prepared. The first step was to state and describe, for each benefit, the
consequence(s) through an LCA perspective of each one of its benefits on soil,
plant, environment, farmer or harvest. After that, the quantification of the substi-
tuted or saved process based on up-to-date knowledge and data was discussed.
Thirdly, the impact categories in existing impact assessment methodologies which
are most affected when considering compost application, were addressed, together
with the current status of new assessment methodologies. Finally, the nine benefits
studied were classified according to the existing evidences for the positive effects on
soil and plant, the possibility of quantification, and the current availability of tools
for their consideration in LCA.
3 Review of Compost Benefits
A review of the literature dealing with the nine potential benefits resulting from
compost application reflected in the scientific literature is presented in the next
sections. For each one, a discussion of the main factors affecting the performance of
the benefit, the degree of proof and the range of the effects measured are included.
3.1 Nutrient Supply
An adequate use of composted products, together with a mineral nitrogen supple-

mentation, can provide the same fertilization level as mineral NPK fertilizers, while
saving the economic and the environmental costs associated with the production
and use of inorganic fertilizers. Moreover, compost can both provide essential trace
minerals to the soil that are not supplied when mineral fertilizers are added (Favoino
and Hogg 2008), and increase the cation exchange capacity (Diacono and
Montemurro 2010). This favours a better retention of nutrients in the soil, thereby
increasing fertilization efficiency.
The content of nutrients in composts is highly variable, depending on the raw
material composition and the composting conditions. For example, Boldrin et al.
(2009) reported that the typical contents of N, P and K in biowaste compost are
7–28, 1–9 and 3–23 kg per ton of dry compost, respectively. However, these nutri-
ents are not immediately available to the growing plant (Favoino and Hogg 2008).
Because of low nutrient availability and slow mineralization rates, some authors
claim that compost is not a good fertilization supplier (Fitzpatrick 2004; Moral and
Muro 2008), although it is complementarily enhancing physical and biological fer-
tility of soils (Gosling and Rayns 2008). In fact, a period of net immobilization of N
after compost application was reported for some case studies, and an extra applica-
tion of mineral N was required to prevent N deficiency. Initial N-immobilisation is
less likely with mature compost with a low C/N ratio, because N-immobilization
already occurred during the composting process (Amlinger et al. 2003b; Bar-Tal
et al. 2004; Hargreaves et al. 2008; ROU 2007).
The fate of N after compost application depends in general on the compost qual-
ity, the nutrient content of raw material, management of composting process, etc.,
together with site and management Factors such as climatic conditions and period
of the year, time of application, soil properties, mechanical aeration, crop rotation,
and plant-soil interactions.
Based on the literature (Table 1), it could be stated that between 5% and 22% of
the N contained in compost is available the first year, and 40–50% in 3–5 years.
However, most of these studies have been done for temperate climates, while in
warmest climates the mineralization rates can increase (Sikora and Szmidt 2004).
Accordingly, two studies located in the Mediterranean region and including experi-
mental measurement of the mineralization rates, are reporting rates of 15–24% in
4–5 months (Hadas and Portnoy 1997; Elherradi et al. 2005). Rates for P and K
262
Table 1 Literature review of nutrient released from compost to soils during 1–5 years after its application. Between 5% and 60% of the applied with compost
N is mineralized, depending on the time frame considered. Figures range between 35% and 100% for P and between 75% and 100% for K
Compost N released from total (%) P released from total (%) K released from total (%)
Study feedstock Short-term Mid-term Short-term Mid-term Short-term Mid-term
Reviews
Amlinger et al. Biowaste and 5–15% (1 yr) 2–8% (each yr)
(2003b) garden waste 50% (5 yr)
Amlinger et al. Biowaste 10–15% (1 yr) 35% (1 yr) 75% (1 yr)
(2003a)
ROU (2007) Food and garden 15% (1 yr) 40% (3–5 yr) 38% (1 yr) 100% (3–5 yr) 80% (1 yr) 100% (3–5 yr)
waste
Hargreaves MSW 16–21% (6 months)
et al.(2008) 10% (1 yr)
10–22% (1 yr)
Diacono and Several feedstocks 15–20 (1 yr) 3–8% (each yr)
Montemurro
(2010)
Individual papers
Terman et al. MSW 16%
(1973)a
Murillo et al. MSW 22%
(1995)a
Hadas and MSW 22% (5 months)
Portnoy
(1997)a
J. Martínez-Blanco et al.
Biala and Wynen Garden waste 15% (1 yr) 25–35% (2 yr) 50% (1 yr) 50% (2 yr) 80% (1 yr) 20% (2 yr)
(1998)a
Sikora (1997)a MSW 10%
Sullivan et al. Food waste 7.6–8.1%
(1998)a
Mamo et al. MSW 0–12% (1 yr)
(1999)a
Frossard et al. Organic solid 2–16% 40–77%
(2002)a waste
Houot et al. MSW 3%
(2002)a
Sullivan et al. Food waste 0%
(2002)a
Guerini et al. MSW 20% (1 yr) 10% (2 yr)
(2006) 5% (3 yr)
Hansen et al. MSW 30% (1 yr) 30%
(2006) 30% (2 yr)
Elherradi et al. Household 15–24% (4 months)
(2005)
MSW municipal solid waste, yr year
a
Reviewed in ROU (2007)
Assessing the Environmental Benefits of Compost Use-on-Land…
263
reported in the literature are shown in Table 1. Boldrin et al. (2009) reported that
final – i.e. long-term – utilization efficiencies for N, P and K were in the order of
20–60% for N, 90–100% for P, and 100% for K, without taking into consideration
the excess of P and K with respect to the N applied.
3.2 Carbon Sequestration
The SOC pool is the largest terrestrial reservoir of organic C (Lal 2004a). More than
twice as much carbon is held in soils as in vegetation or in the atmosphere (Whalen
and Sampedro 2010). Pools of SOC are defined, as active, slow, and resistant accord-
ing to their decomposition rate (Paul et al. 2001). Increasing SOC stocks is one of
the strategies with the largest potential for climate change mitigation (Rogner et al.
2007). The fate of SOC stocks is largely determined by the land management, as for
example intensive agricultural practices can deplete the SOM and result in a net
release of SOC to the atmosphere.
The use of compost could represent an effective alternative for increasing SOC
stocks (Lal 2004a). Table 2 shows some examples of compost effects on soil carbon
sequestration. For example, Fortuna et al. (2003) reported that application of compost
for 4 years to a crop rotation increased the slow pool C by 75% and the resistant SOC
pool by 40%. Similarly, Lal (2002) estimated that the use of 10–20 t ha−1 of compost
per year has the potential to increase SOC between 500 and 1,000 kg ha1 year−1.
In general terms, sequestration rates are highly variable and depend on several
compost, and site-dependent characteristics. Sequestration potentials are higher
with high application rates and with mature composts (Eghball 2002), as the labile
C fractions are rapidly metabolized and only the resistant ones remain stable in the
soil (Fabrizio et al. 2009; Sodhi et al. 2009a). With regards to the soil characteristics,
an important factor is the limited capacity of the soil to act as a C sink: SOC content
increases rapidly when organic amendment is added to soils where SOM is depleted,
while the increase rate is slower when the SOC content is close to saturation i.e. the
maximum amount of stable C that one soil can maintain (Stewart et al. 2007). It can
take over 50 years of continuous restorative practices to reach this new equilibrium
in SOC level, although much of the change takes place in the first 10 years and
slow down when soils are close to a saturation level (Audsley et al. 2003). Finally,
climate conditions determine mineralization rates and the residence time of SOC in
soil, as mineralization rates tend to be higher in warm climates (Sikora and Szmidt
2004).
Time horizon chosen also plays an important role in the calculation of the seques-
tration potential. For short-term field studies between 40% and 70% of the C applied,
remained in the soil shortly after compost incorporation, i.e. 3–6 months (Fabrizio
et al. 2009; Fagnano et al. 2011). On a much longer time-horizon, more relevant for
the Intergovermnental Panel on Climate Change (IPCC) and LCA approach, other
studies estimated that between 2% and 14% of the carbon introduced with a single
compost application is still bound to soil after 100 years (Boldrin et al. 2009;
Table 2 Literature review of compost effects on soil carbon sequestration. Carbon sequestration rates have shown to be higher in the short term (up to 40% of
the applied C) decreasing over time to 16%
Dosage ww
Study Compost feedstock (t ha−1 yr−1) C addition Crop Type of soil Length (yr) C retained (%)
a
Smith et al. (2001) 100 8.2
Eghball (2002) Feedlot manure 21.5b 1.7 t ha−1 yr−1 Maize Silty clayy loam, 4 36
mesic typic
argiudolls
Fortuna et al. Oak leaves-manure (1:1) 4.48b Maize-wheat Kalamazoo loam 4 63
(2003)
Sánchez-Monedero Two phase olive mill 48 25.44 t ha−1 yr−1 No crop na 0.77 63.1
et al. (2008)c waste + sheep litter
Two phase olive mill 48 25.48 t ha−1 yr−1 No crop na 0.77 66.1
waste + sheep
litter + grape stalks
waste + sheep litter
waste + sheep
litter + grape stalks
Sodhi et al. Rice straw compost 4 1.24 t ha−1 Rice–wheat Sandy-loam typic 10 29
(2009b) ustipsamment
Rice straw compost 16 4.96 t ha−1 Rice–wheat Sandy-loam typic 10 29
ustipsamment
Fabrizio et al. Food residues and 50 2.72 g kg−1 soil Maize Silt-clay fluvic- 0.41 40
(2009) ligno-cellulosic eutric cambisol
wastes
Food residues and 85 4.64 g kg−1 soil Maize Silt-clay fluvic- 0.41 53
ligno-cellulosic eutric cambisol
wastes
(continued)
265
Table 2 (continued)
266
Dosage ww
Study Compost feedstock (t ha−1 yr−1) C addition Crop Type of soil Length (yr) C retained (%)
Fagnano et al. MSW 10 1,282 kg ha−1 Lettuce Sandy loam eutric 0.5 63.1
(2011) regosol
MSW 30 3,847 kg ha−1 Lettuce Sandy loam eutric 0.5 70.3
regosol
MSW 60 7,694 kg ha−1 Lettuce Sandy loam eutric 0.5 69.9
regosol
MSW municipal solid waste, yr year, na not available
a
Modeling study
b
Dry weight
c
Incubation study
Favoino and Hogg 2008; Smith et al. 2001). To include all the above mentioned fac-
tors, quantification of the exact amount of C retained in soil after compost applica-
tion may thus need to be assessed through the use of complex agroecosystem models
that incorporate mineralization dynamics for the case-specific conditions (Bruun
et al. 2006).
3.3 Weed, Pest and Disease Suppression
Weeds reduce crop quality and yield by competing for light, water and nutrients.
Organic composted mulches could effectively control weeds in crops, thus avoiding
herbicide applications totally or partially, while compost does not have relevant
effects on weed grow and density when used as a soil amendment (De Cauwer et al.
2010; ROU 2007).
Together with weeds, the incidence of pest and disease is one of the major factors
limiting the productivity of agro-ecosystems. Modern agriculture heavily relies on
the use of synthetic pesticides, although they are proved to damage the environment
and human health (Streck 2003) and they often have unsatisfactory effectiveness
(Bonanomi et al. 2007; Litterick et al. 2004). Litterick et al. (2004) pointed out sev-
eral integrated crop protection strategies as potential alternatives to the use of syn-
thetic pesticides, including crop rotation, local resistant varieties, biological control
agents, and, as it is explained below, suppressive compost application.
Several factors concur for a successful biological control of plant pathogens with
composts. During the composting process, the use of heterogeneous input material(s),
a sufficient and controlled increase in temperature, aeration and moisture, a correct
maturation process, and non-anaerobic storage are the most important recommenda-
tions to obtain a suppressive compost (Bonanomi et al. 2007, 2010; De Bertoldi 2010;
Litterick et al. 2004; ROU 2007). Microbial diversity and the presence of specific sup-
pressive micro-organisms or consortia in compost are also suggested to be essential
for disease suppression (Amlinger et al. 2003a; De Bertoldi 2010; Termorshuizen
et al. 2006). High compost application rates are related to higher suppression and a lag
of time between compost application and planting is advisable (Amlinger et al. 2003a).
The disease suppressive properties of composts are induced during the curing phase,
because most biocontrol agents, mainly fungal and bacterial species, re-colonize the
compost from the surrounding air and soil after peak heating (De Bertoldi 2010;
Litterick et al. 2004; ROU 2007). However, several authors argued that excessively
stabilized organic matter may not support adequate activity of biocontrol agents due
to the scarcity of available nutrients (ROU 2007; Bonanomi et al. 2010).
A decreased incidence of diseases after compost application to soil could be attri-
buted either to a direct suppression of the pathogens or to a systemic resistance induced
in the crop (Noble and Coventry 2005). Nevertheless, recent publications indicated that
there is still insufficient knowledge about the general principles of disease suppression
by compost, because it is often pathogen-specific and related to the mechanism(s) of
disease suppression (Bonanomi et al. 2010; Termorshuizen et al. 2006).
Fig. 3 Effect of compost amendments on disease incidence and severity caused by soilborne
pathogens, compared to the non-amended control. Data are the percentage of cases with highly
suppressive (>80% disease reduction), suppressive (significant disease reduction), null (no
significant effect) or conducive effects (significant disease increase). The total suppressive cases
are the sum of highly suppressive and suppressive. Only combinations with at least ten studies are
shown (Drawn from Bonanomi et al. 2007)
The application of composts can rarely – if ever – result in a disease control

comparable to the use of chemicals (Litterick et al. 2004). Bonanomi et al. (2007)
reviewed 250 studies about direct suppression of soil-borne fungal diseases in pot
and field experiments, reporting relevant suppressive effects for several types of
compost in 58–74% of the studied cases for Phytium spp, Fusarium spp, Phytophthora
spp and Verticillium dahlia (Fig. 3). For the other pathogens the suppression was
lower or null. However, only 12% of the cases studied resulted in a disease reduction
>80% when organic amendments were applied, which is the minimum pathogen
population reduction that farmers are using as a criterion for replacing chemical
pesticides (Bonanomi et al. 2007). Precise information about the duration of the sup-
pressive effects was not found; however, most of the reviewed studies estimated the
suppression was efficient for 3–6 months after the application of the amendment.
It should be mentioned that there is also a risk of pathogens and unwanted plant
seeds being present in compost (Vinneras et al. 2010). Nevertheless, this risk can be
minimized with a proper combination of temperature (64–70°C), processing dura-
tion (21 days), and turning frequency during composting (De Cauwer et al. 2010;
Hargreaves et al. 2008; Noble and Roberts 2004).
In conclusion, the extent to which pest and diseases are suppressed when com-
post is used cannot be univocally determined, mainly because of lack of evidence in
field conditions, as well as because the rate of suppression is closely dependant on
the type of pathogen and the interrelation with the analysed plant. While there is a
considerable amount of data covering pot experiments, field experiments lack and/
or published studies only focused on soil-borne diseases. Results from laboratory
experiments cannot be directly extrapolated to field crops, because field conditions
are much more variable (Fuchs 2010) and because much smaller quantities of
compost are typically used in the field compared to pot conditions.
3.4 Crop Yield
As the cost of fertilizers is often small compared to the cost of lost yield, farmers
prefer to over-fertilize their crops with N rather than risking to under-fertilize with
the consequent loss of revenue (Del Amor 2007). Moreover, agricultural producers
often claim that crop yields are much lower with organic fertilization than with
inorganic fertilizers (Mäder et al. 2002). However, excess nitrogen may result in
lodging, greater weed competition and pest attacks, with substantial losses of pro-
duction. In addition, the nitrogen not taken up by the crop is likely to be lost to the
environment, which potentially contributes to groundwater and atmospheric pollu-
tion (FAO 1998; Roy et al. 2006). Sustainable agriculture would ideally produce
good crop yields with minimal impact on the ecological system (Mäder et al. 2002).
Optimized use of fertilizers should be targeted upon several factors, including the
soil nutrient pool, soil management, compost composition and maturity, rates and
methods of application, crop species and variety, fertiliser rates, and intervals
between application and planting (Amlinger et al. 2003b; Diacono and Montemurro
2010; ROU 2007).
Application of MSW compost had positive effects on growth and yield of a wide
variety of crops (Shiralipour et al. 1992). Experimental tests show that trials amended
with biowaste compost provide higher yields than controls not receiving any treatment
(Table 3). However, when compost-based fertilization is compared to mineral fertil-
ization, no relevant differences in crop yields are reported: non-significant differences
were observed in 64% of the cases; while lower and higher yields for the composting
option were seen in 32% and 4% of the cases respectively (Table 3). The best agro-
nomic performance of compost is obtained with high dosages and high frequency of
application, and by suplementing compost with mineral fertilizers to achieve a bal-
anced supply of nutrients (Amlinger et al. 2003a; Diacono and Montemurro 2010;
ROU 2007). In general, suplementing compost with mineral fertilizers leads to yields
similar to the conventional production pathway (Table 3). When compost is used
alone, long-term and repeated applications are needed to achieve a steady state that
guarantees a crop yield close to the inorganic fertilizing scheme. Yield increases
in fields transitioning from conventional to organic production systems usually
require 3–5 years to be detected (Herencia et al. 2008). In conclusion, compost effects
on crop yields are variable ranging from decreases of 495% to increases of 52%.
Table 3 Literature review of crop yield effects after compost application in soils. Impacts in crop yields vary from decreases of 495% to increases of 52%
270
Supplementary
Compost Dosages ww fertilizing Effect on Nutrient Minerali-
Study feedstock (t ha−1 yr−1) treatment Crop Type of soil Length (yr) yield (%) balance zation rate
Comparison with mineral fertilizer treatment
Iglesias- MSW 10 no Ryegrass Andeptic 0.5 M>C yes nm
Jimenez paleudult (+136%)
and Alvarez MSW 50 no Ryegrass Andeptic 0.5 M>C yes nm
(1993)a paleudult (+495%)
CIWMB Grass 13b NPK Cotton + winter Sandy loam 1 ns no no
(1997) clippings forage crop
Grass 15b NPK Irrigated corn/ sandy loam 1 M>C no no
clippings wheat (+8%)
Diez et al. MOW 28c 150 Kg N ha−1 Maize Sandy-loam 3 ns nm yes
(1997) MOW 28c no Wheat Sandy-loam 3 ns nm yes
MOW 28c no Maize Sandy-loam 3 ns nm yes
Bazzoffi et al. MSW 96c no Maize Clay loam typic 1 ns no no
(1998) udorthent
MSW 96c no maize Clay loam typic 2 ns no no
udorthent
MSW 96c no Maize Clay loam typic 3 M>C no no
udorthent (+71%)
Eriksen et al. MSW 63c no Maize Sandy, 2 ns nm no
(1999) siliceous,
mesic
psammentic
hapludults
MSW 126c no Maize Sandy, siliceous, 2 ns nm no
mesic
psammentic
hapludults
Supplementary
MSW 189c no Maize Sandy, 2 ns nm no
siliceous,
mesic
psammentic
hapludults
Mamo et al. MSW 90 125 Kg N ha−1 Corn Loam sandy 3 ns no no
(1999) MSW 90 250 Kg N ha−1 Corn Loam sandy 3 ns no no
Wolkowski MSW 35b no Grain Saybrook silt 1 M>C nm nm
(2003) loam (+37%)
MSW 138b no Grain Saybrook silt 1 M>C nm nm
loam (+28%)
MSW 35b no Grain Boyer loamy 1 M<C nm nm
sand (−22%)
MSW 138b no Grain Boyer loamy 1 M<C nm nm
sand (−52%)
MSW 35b no Grain Saybrook silt 2 M>C nm nm
loam (+11%)
MSW 138b no Grain Saybrook silt 2 ns nm nm
loam
MSW 35b no Grain Boyer loamy 2 ns nm nm
sand
MSW 138b no Grain Boyer loamy 2 ns nm nm

sand
Elherradi et al. Household 10 no Lettuce Sandy 0.3 M>C no no
(2005)a (+98%)
Household 10 no Lettuce Loamy-clay 0.3 M>C no no
(+39%)
Household 20 no Lettuce Sandy 0.3 M>C no no
(+81%)
Household 20 no Lettuce Loamy-clay 0.3 ns no no
(continued)
271
272
Table 3 (continued)
Supplementary
Household 30 no Lettuce Sandy 0.3 M>C no no
(+45%)
Household 30 no Lettuce Loamy-clay 0.3 ns no no
Household 10 50% NPK Lettuce Sandy 0.3 M>C no no
(+37%)
Household 10 50% NPK Lettuce Loamy-clay 0.3 ns no no
Household 20 50% NPK Lettuce Sandy 0.3 M>C no no
(+27%)
Household 30 50% NPK Lettuce Sandy 0.3 ns no no
Mkhabela and MSW 17 no Potato Pugwash sandy 1 M>C no no
Warman loam (+44%)
(2005)d
MSW 33 no Potato Pugwash sandy 1 M>C no no
loam (+50%)
MSW 50 no Potato Pugwash sandy 1 ns no no
loam
loam
Supplementary
loam
loam
MSW 6 no Sweet corn Pugwash sandy 1 ns no no
loam
MSW 12 no Sweet corn Pugwash sandy 1 M>C no no
loam (+18%)
MSW 17 no Sweet corn Pugwash sandy 1 M>C no no
loam (+24%)
loam
loam
loam
Montemurro MSW 5 50% N Sunflower Silty-clay 2 ns yes no
et al. (2005) MSW 9 no Sunflower Silty-clay 2 ns yes no
Guerini et al. MSW 60 NPK Maize Sandy loam 5 ns yes yes
(2006) MSW 60 NPK Maize Clay loam 6 ns yes yes
Montemurro MSW na Alfalfa 3 ns yes nm
et al. (2006) MSW na Cocksfoot 3 M>C yes nm

(+21%)
Ros et al. Urban organic 14.6 no Maize Loamy silt 12 M>C nm yes
(2006a) waste (+23%)
Urban organic 14.6 80 kg N ha−1 Maize Loamy silt 12 ns nm yes
waste
Green waste 10.9 no Maize Loamy silt 12 M>C nm yes
(+16%)
(continued)
273
274
Table 3 (continued)
Supplementary
Green waste 10.9 80 kg N ha−1 Maize Loamy silt 12 ns nm yes
Ghorbani et al. Household 20c no Tomato Sandy clay 1 ns nm no
(2008) loam
Household 20c no Tomato Sandy clay 2 M>C nm no
loam (+33%)
Odlare et al. Separated 4b 50 kg N ha−1 Oat + spring Sandy-clay 4 ns no yes
(2008) household barley loam Eutric
waste + gar- cambisol
den litter
Separated 8b no Oat + spring Sandy-clay 4 M>C no yes
household barley loam Eutric (+44%)
waste + gar- cambisol
den litter
Coria-Cayupan Fruit and 10 no Lettuce nm Crop ns yes nm
et al. (2009) vegetables period
MSW 10 no Lettuce nm crop period M>C yes nm
(+138%)
Montemurro MSW 5 50% N Winter wheat Silty-clay 4 M<C yes no
(2009) (−7%)
MSW 9 no Winter wheat Silty-clay 4 M>C yes no
(+11%)
Celik et al. Grass, wheat 38b no Winter wheat Typic 13 M>C no no
(2010) stubbles xerofluvent (+56%)
and plant clay–loam
leaves
Supplementary
Morra et al. MSW 15 no Horticulture Sandy loam 3 ns no no
(2010) rotation calcaric
cambisol
MSW 30 no Horticulture Sandy loam 3 ns no no
rotation calcaric
cambisol
MSW 45 no Horticulture Sandy loam 3 ns no no
rotation calcaric
cambisol
MSW 15 50% NPK Horticulture Sandy loam 3 ns no no
rotation calcaric
cambisol
MSW 15 25% NPK Horticulture Sandy loam 3 ns no no
rotation calcaric
cambisol
Comparison with control treatment
Iglesias- MSW 10.0 no Ryegrass Andeptic 0.5 N<C
Jimenez paleudult (+30%)
and Alvarez MSW 50.0 no Ryegrass Andeptic 0.5 N<C
(1993)a paleudult (+106%)
Sabrah et al. Urban waste 16.5 NPK Irrigated wheat Sandy 1 ns
(1995)
Urban waste 16.5 NPK Irrigated wheat Sandy 2 ns

Urban waste 33 NPK Irrigated wheat Sandy 1 N<C
(+11%)
(+34%)
Urban waste 49.5 NPK Irrigated wheat Sandy 1 N<C
(+36%)
(continued)
275
276
Table 3 (continued)
Supplementary
Urban waste 49.5 NPK Irrigated wheat Sandy 2 N<C
(+43%)
(+45%)
(+48%)
Mamo et al. MSW 90.0 no Corn Loam sandy 3 N<C
(1999) (+71%)
Movahedi Urban 50.0 125 Kg N ha−1 Rainfed maize Silt loam 1 N<C
Naeini and compost (+10%)
Cook Urban 50.0 125 Kg N ha−1 Rainfed maize silt loam 1 ns
(2000) compost
Hartl et al. Biowaste 22d no Rye Calcaric 5 ns
(2003) fluvisol
Biowaste 12d no Rye Calcaric 5 ns
fluvisol
Tejada et al. Crushed cotton 16 no Rice Aquic 3 N < C (+5%)
(2006a) gyn xerofluvent
Crushed cotton 24 no Rice Aquic 3 N < C (+7%)
gyn xerofluvent
Supplementary
Crushed cotton 32 no Rice Aquic 3 N < C (+7%)
gyn xerofluvent
Crushed cotton 16 250 kg N Rice Aquic 3 N < C (+7%)
gyn xerofluvent
Crushed cotton 24 250 kg N Rice Aquic 3 N < C (+9%)
gyn xerofluvent
Crushed cotton 32 250 kg N Rice Aquic 3 N<C
gyn xerofluvent (+11%)
MSW municipal solid waste, MOW municipal organic waste, M mineral fertilizers option, C compost option, N option of no fertilization, na not available, nm
not mentioned, ns no significant differences
A > B, A is higher than B; A < B, A is lower than B
a
Case study of pot experiment
b
Only dry weight is available in the study. We considered a 35% moisture content for the ww calculation
c
Compost was applied once at the beginning of the experiment
d
The dosages were different each year. An average value is provided
277
Impacts depend largely on the application rates, number of applications over time
and management factors such as supplementing with inorganic fertilizers.
3.5 Soil Erosion
Soil erosion involves great economic losses and environmental damage due to the
decline in soil productivity and soil functions and, in the worst scenarios, the irrevers-
ible loss of cultivable land. Indirect effects associated to soil erosion are the contamina-
tion of streams with soil particles and agrochemicals, the incidence of respiratory
diseases in connection with particles transported by wind, and the associated loss of
SOM, soil nutrients and beneficial agricultural microorganisms (Gardner et al. 2012).
Organic matter has a key role in stabilizing soil structure, as it increases the inter-
particle cohesion within aggregates and enhances the aggregate’s hydrophobicity, thus
decreasing their breakdown (Diacono and Montemurro 2010; Ruehlmann and Körschens
2009; Soane 1990). Aggregate stability determines soil sensitivity to crusting and ero-
sion (Bissonnais 1996). The application of easily decomposable organic substances has
an intense and transient effect on aggregate stability while more recalcitrant substances
have a lower but longer term effect (Abiven et al. 2009). Several studies showed that
compost addition significantly increases soil aggregate stability therefore making it
more resistant to erosion. Short-term incorporation of biowaste compost to soil can
increase aggregate stability as much as 41% and decrease aggregate instability between
6.9% and 30% depending on the dosage (Table 4). Short-term experiments under simu-
lated rain showed that soil runoff can be reduced by 66% with compost incorporation.
Data concerning mid- and long-term effects of biowaste compost on soil erosion are
scarce, and thus data regarding other types of compost were also included (Table 4).
Large variability in compost effects was observed in mid-term experiments, with
increases in aggregate stability ranging from 0% to 63% and decreases in aggregate
instability of 21% (Table 4). As a consequence, soil loss was reduced between 5% and
36%, and soil runoff between 26% and 30% (Table 4). With regards to long-term com-
post application, i.e. 10 years, Sodhi et al. (2009a) reported that rice straw compost
application to soil may increase aggregate stability by 4%. No other studies covering
long-term perspective could be found. Finally, it should be noted that field studies were
carried out in Mediterranean areas, while data from other regions is scarce.
3.6 Soil Moisture Content
Agriculture is by far the largest water-using sector, accounting for about 70% of the
consumed freshwater coming from rivers and aquifers (FAO 2008). Several mea-
sures are proposed for reducing water demand in the agricultural sector, including
efficient irrigation, reduction of distribution losses, maximisation of effluent reuse,
and conservation of soil moisture.
The amount of soil water available to plants is determined by the depth of the
roots and the moisture storage capacity of the soil. Storage capacity is linked to
Table 4 Literature review of the effect of compost application on the soil parameters related to soil erosion. Compost showed to increase soil aggregate
stability between 29% and 63%, reducing soil loss between 5% and 36%
Dosage ww Length Change (%) after
Study Compost feedstock (t ha−1 year−1) Crop Type of soil (year) compost application
% Water stable aggregates
Sodhi et al. (2009b) Rice straw 8 Rice-wheat Typic ustipsamment 10 4
Soil structural stability or aggregate stability
Tejada and González (2006b)a Cotton gin crushed 30 Wheat Typic xerofluvent 5 0
Annabi et al. (2006)b MSW 5 29.3
Annabi et al. (2007)c MSW 34.59 g kg−1 No crop Typic hapludalf <1 41
Biowaste 74.79 g kg−1 No crop Typic hapludalf <1 29
Green waste + sludge 21.3 g kg−1 No crop Typic hapludalf <1 29
Bipfubusa et al. (2008)d Paper sludge 40 Corn Silt loam 4 45
Leroy et al. (2008) Vegetable, fruit and 22.5 Corn Sandy loam 9 63
garden waste
Tejada et al. (2008) Green manure + beet 10 No crop Xelloric calciorthid 4 10.5
vinase
Tejada et al. (2009b) Plant residues (rapeseed) 17.7 No crop Xelloric calciorthid 4 28.3
Arthur et al. (2011) Garden waste 30 m3 ha−1 Cauliflower Sandy loam, haplic 9 45
podzol
Vegetable, fruit and 30 m3 ha−1 Cauliflower Sandy loam, haplic 9 27.5

garden waste podzol
Spent mushroom 30 m3 ha−1 Cauliflower Sandy loam, haplic 9 25
podzol
Aggregate instability
Aggelides and Londra (2000) MSW + sewage 39 Grass fallow Clay soil, humic 1 −6.89
sludge + saw dust fluvaquent
(continued)
279
280
Table 4 (continued)
MSW + sewage 78 Grass fallow Clay soil, humic 1 −6.89
MSW + sewage 156 Grass fallow Clay soil, humic 1 −17
MSW + sewage 39 Grass fallow Loam soil, typic 1 −13
sludge + saw dust xerochrept
Tejada and González (2008)a Cotton gin crushed 20 No crop Xerollic calciorthid 4 −21
Soil loss
Bazzoffi et al. (1998)e 64 Corn Clay loam, typic 3 −5
udorthent,
calcaric regosol
Tejada and González (2006b)a Cotton gin crushed 30 Wheat Typic xerofluvent 5 −36
Tejada and González (2008)b Cotton gin crushed 20 No crop Xerollic calciorthid 4 −29.2
Cotton gin crushed 20 No crop Xerollic calciorthid 4 −29.2
Tejada et al. (2009a)a Beet vinasse + greenwaste 15.82 No crop Xerollic calciorthid 3 −28.9
vermicompost
Runnoff
Bresson et al. (2001)a, c MSW 50 No crop Silt loam, 60 min −66.5
typic hapludalf
Arthur et al. (2011) Garden waste 30 m3 ha−1 Cauliflower Sandy loam, haplic 9 −26
podzol
Vegetable, fruit and 30 m3 ha−1 Cauliflower Sandy loam, haplic 9 −30
garden waste podzol
Spent mushroom 30 m3 ha−1 Cauliflower Sandy loam, haplic 9 −30
podzol
Soil erodibility
Arthur et al. (2011) Garden waste 30 m3 ha−1 Cauliflower Sandy loam, haplic 9 −18
podzol
Vegetable, fruit and 30 m3 ha−1 Cauliflower Sandy loam, haplic 9 −27
garden waste podzol
Spent mushroom 30 m3 ha−1 Cauliflower Sandy loam, haplic 9 −18
podzol
a
Data obtained with simulated rain
b
Three applications over 5 years
c
Incubation study
d
Three applications per year during 2 years
e
Only one compost application at the beginning of the experiment
281
porosity, and thus the texture and structure are the critical factors (Rawls et al. 2003),
meaning that poor structure, low SOM, low carbonate content and presence of stones
reduce the moisture storage capacity of a given soil texture class. The addition of
organic matter increases the number of micropores and macropores (Bot and Benites
2005). As a consequence, less irrigation water is needed to irrigate the same crop.
Compost has a high water holding capacity because of its organic matter content,
which in turn improves the water holding capacity of the soil (Hargreaves et al.
2008). Large increases in water holding capacity are reported for high-rate compost
application on soils with initially low SOC content (Glab et al. 2009; Olness and
Archer 2005; Rawls et al. 2003; ROU 2007). Additionally, the application of com-
posted organic matter reduces surface sealing, improves infiltration, lowers the
water table while maintaining the same moisture, and reduces runoff generation
(Pandey and Shukla 2006; ROU 2007).
Olness and Archer (2005) applied the General Energy Model for Limited Systems
to U.S. National Soil Inventory Database and indicated that 1% increase in SOC
causes a 2–5% increase in water holding capacity depending on the soil texture. In
other studies, the same authors reported increases in water holding capacity ranging
from about 0.8% to 8.4% for each percent increase in SOC. Accordingly, Hudson
(1994) showed that for each 1% increase in SOM, the available water holding capac-
ity in the soil increased by 2.2–3.7%.
With regards to application of biowaste compost, the reviewed literature (Table 5)
showed that when compost is applied at low rates (< 16 t ha−1 year−1) no effects are
observed in the moisture content in soils, even if compost application continues for
several years (Castillejo and Castello 2010; CIWMB 1997; Glab et al. 2009;
Hortenstine and Rothwell 1973; Suzuki et al. 2007). Sabrah et al. (1995) measured
an increase on plant available water, the portion of the water holding capacity that
can be absorbed by a plant without stress problems, from 16.5% to 43% when
16.5–66 t ha−1 year−1 of compost were applied. Movahedi Naeini and Cook (2000)
measured less than 2% of increase in water holding capacity after the application of
50 t ha−1 of compost in a silt loam soil. Discordant results are reported for sandy
soils; for example, Hortenstine and Rothwell (1973) found that applications of
compost ranging 32–128 t ha−1 year−1 led to water holding capacity proportional
increases of 5–50%, whereas Weber et al. (2007) did not detect any difference in
WHC after 1 year of 120 t ha−1 application. However, even in soils where compost
does not increase PAW per unit of soil volume, compost amendments increased the
availability of water to plants by facilitating denser and deeper root growth, Fig. 4
(Curtis and Claassen 2005).
3.7 Soil Workability
Soil workability is a measure of how easy it is to till or plough a soil. Workability

depends on several interrelated soil properties, including soil texture, SOM content,
structure, bulk density, and presence of gravel and stones (Fischer et al. 2008).
Intensive agricultural practices may lead to significant losses of the SOM increasing
Table 5 Literature review of soil moisture content after compost application in soils. Water holding capacity and plant available water can increase by 50 by
34% respectively
Change on the
Compost Dosages ww water capacity
Study feedstock (t·ha−1·yr−1) Crop Type of soil Length (yr) indicator (%)
Water savings
CIWMB (1997) Grass clippings 13.3a Irrigated corn/wheat Sandy loam 1 0
Grass clippings 14.9a Irrigated corn/wheat Sandy loam 2 0
Glab et al. (2009) Biowaste 7 Rotation cereal/ Mollic-gleyic fluvisol 13 0
potato
Biowaste 13 Rotation cereal/ Mollic-gleyic fluvisol 13 0
potato
Biowaste 18 Rotation cereal/ Mollic-gleyic fluvisol 13 0
potato
Water holding capacity (WHC)
Hortenstine and Municipal waste 16 Irrigated sorghum Sandy na 0
Rothwell (1973) Municipal waste 32 Irrigated sorghum Sandy na 5
Municipal waste 64 Irrigated sorghum Sandy na 26
Municipal waste 128 Irrigated sorghum Sandy na 50
Mamo et al. (1999) Urban waste 138.5a Irrigated corn Loamy sand na 4.4
Movahedi Naeini and Urban waste 50 Rainfed corn Silt loam 1 2

Cook (2000) Urban waste 50 Rainfed corn Silt loam 1 0
Bazzoffi et al. (2006)b MSW 60c No crop Silty clay soil typic 1 7
eutrochrept
MSW 60c No crop Silty clay soil typic 3 8
eutrochrept
eutrochrept
eutrochrept
283
(continued)
284
Table 5 (continued)
Change on the
c
MSW 60 No crop Sandy loam texture 1 0
typic udifluvent
MSW 60c No crop Sandy loam texture 3 0
typic udifluvent
typic udifluvent
typic udifluvent
Suzuki et al. (2007) Leaf litter 10 Forage sorghum Sandy soil 1 0
Albaladejo et al. (2009) MSW 205.5 No crop Typic calcisol 1 18
MSW 342.5 No crop Typic calcisol 1 26
Castillejo and Castelló MSW 10 Fodder shrub Gypsum spoil (soil 1 0
(2010)d (Atriplex halimus) forming material)
MSW 30 Fodder shrub Gypsum spoil (soil 1 19
(Atriplex halimus) forming material)
MSW 50 Fodder shrub Gypsum spoil (soil 1 25
(Atriplex halimus) forming material)
Plant available water (PAW)
Sabrah et al. (1995) Urban waste 16.5 Irrigated wheat Sandy 2 16.5
Urban waste 33 Irrigated wheat Sandy 2 30
Urban waste 49.5 Irrigated wheat Sandy 2 35
Urban waste 66.0 Irrigated wheat Sandy 2 43

Change on the
a
Curtis and Claassen Garden waste 415.4 Grass (Elymus Serpentinitic, thermic 1.5 0
(2005)d elymoides) lithic argixerolls
Garden waste 830.8a Grass (Elymus Serpentinitic, thermic 1.5 126
elymoides) lithic argixerolls
Curtis and Claassen Yard waste 830.8a Big squirreltail Sandy loam Lahar 0.5 −19
(2009)d, c Yard waste 830.8a Needle grass Sandy loam 0.5 −21
serpentinistic
Yard waste 830.8a Needle grass Sandy loam sandstone 0.5 32
Weber et al. (2007) Municipal solid 120,0 No crop Sandy 1 0
waste
MSW municipal solid waste, na not available
a
Only dry weight is available in the study. We considered 35% moisture content for the ww calculation
b
Pot experiment
c
Non-tillage is compared with tillage and compost option. Therefore we are not only comparing compost effects
d
Case study of land reclamation
285
Fig. 4 Moisture content

capacity is generally
increased when high dosages
of compost are applied to
soil. Soil humidity is
measured with a tensiometer
in this horticultural plot
(Photo by Pere Muñoz)
soil bulk density and compaction while decreasing workability. Soil compaction is
directly related to reduced yield and quality of crops, increased erosion, and
increased power requirements for tillage (Soane 1990).
Several studies have shown that organic matter addition to the soil decreases bulk
density, and increases aggregate stability (Ruehlmann and Körschens 2009; Soane
1990) thereby reducing soil compaction. Addition of compost to soil supplies
organic matter and can potentially decrease soil bulk density and soil compaction,
increasing soil workability. In the short term, i.e. <1 year, the application of differ-
ent types of biowaste compost decreased soil bulk density between 2.5% and 21%
as compared to soil without compost (Table 6), while in the mid-term, i.e. 1–10
year, the decrease may be as much as 20.6% and 23.1%, with a minimum decrease
of 0.7% (Table 6). Long-term (13 year) compost application can decrease soil bulk
density by 20.6% (Table 6). Compost application seems to be more effective in the
short term (Weber et al. 2007) and with increasing dosages (Hemmat et al. 2010), as
larger doses produce larger decreases in bulk density.
3.8 Soil Biological Properties and Biodiversity
Farming systems have a strong impact on soil biodiversity. In particular, organic

farming systems fostering the use of organic fertilizer showed to increase soil bio-
diversity significantly (Bengtsson et al. 2005; Garratt et al. 2011; Mäder et al. 2002).
Table 6 Literature review of the effect of compost application on the soil parameters related to soil workability. Soil bulk density is decreased between 0.7%
and 20% after compost application, potentially increasing soil workability
Change in the
Dosage ww workability
Study Compost feedstock (t ha−1 yr−1) Type of soil Crop Length (yr) indicator (%)
Soil bulk density
Martens et al. (1992) 25 2 −10.8
Turner et al. (1994) MSW 134a 2 −15
Illera et al. (1999) MSW 80 1 −13.1
Stamatiadis et al. (1999) Green waste + cow 44 Silt clay loam Broccoli 1 −6.1
manure
Zebarth et al. (1999) 45 4 −15.5
Aggelides and Londra MSW + sewage 75 m3 ha−1 Humic fluvaquent, clay Grass fallow 1 −6.3
(2000) sludge + saw dust soil
MSW + sewage 150 m3 ha−1 Humic fluvaquent, clay Grass fallow 1 −12.5
sludge + saw dust soil
MSW + sewage 75 m3 ha−1 Humic fluvaquent, clay Grass fallow 1 −12
MSW + sewage 150 m3 ha−1 Humic fluvaquent, clay Grass fallow 1 −17

Tejada et al. (2008) Green manure + beet Xelloric calciorthid No crop 4 −13.5
vinase
Tejada and González Cotton gin crushed 7.12 Xerollic calciorthid 4 −19.6
(2008) compost
(continued)
287
288
Table 6 (continued)
Change in the
Courtney and Mullen Municipal sludge 100 nm Barley <1 −8.9
(2008)
Spent mushroom 100 nm Barley <1 −8.9
compost
Diana et al. (2008) Wine-producing 1 Alluvial soil Lettuce <1 −21
residues
Mylavarapu and Zinati 75% MSW + 25% 20 Loamy, siliceous, Parsley 1 −2.5
(2009) biosolids hyperthermic,
grossarenic,
paleudult
Hemmat et al. (2010) MSW 25 Typic haplargids/ Wheat-corn 7 −0.7
calcaric cambisols,
fine-loamy, mixed
MSW 50 Typic haplargids/ Wheat-corn 7 −4.5
calcaric cambisols,
fine-loamy, mixed
MSW 100 Typic haplargids/ Wheat-corn 7 −23.1
Calcaric cambisols,
fine-loamy, mixed
Celik et al. (2010) Green waste 25 Typic xerofluvents, Wheat-corn 13 −20.6
clay loam
Soil porosity
Aggelides and Londra MSW + sewage 75 m3 ha−1 Humic fluvaquent, clay Grass fallow 1 5.4
(2000) sludge + saw dust soil
MSW + sewage 150 m3 ha−1 Humic fluvaquent, clay Grass fallow 1 8.5

Change in the
MSW + sewage 75 m3 ha−1 Humic fluvaquent, clay Grass fallow 1 11
MSW + sewage 150 m3 ha−1 Humic fluvaquent, clay Grass fallow 1 27
Weber et al. (2007) MSW (1) 18 Dystric cambisol, Triticale <1 0.39
sandy
MSW (1) 36 Dystric cambisol, <1 0.24
sandy
sandy
MSW (2) 18 Dystric cambisol, Triticale <1 0.46
sandy
sandy
sandy
MSW municipal solid waste, nm not mentioned
a
Only one compost application

289
In the same way, application of compost to soil has significant effects on above- and
belowground soil biota, both in terms of amount, activity, and diversity of organ-
isms. Griffiths et al. (2010) observed that addition of 2.6 t yr−1 ha−1 of biowaste
compost to a soil during 6 years increased earthworm biomass by 514%, enchytra-
eids by 1100%, collembola by 271%, oribatids by 92%, mesostigmatids by 84%,
microbívorous nematodes by 35% and omnivorous nematodes by 45%.
Due to the central role of microorganisms in soil nutrient cycling, most of the stud-
ies dealing with the impacts of compost on soil biodiversity are focused on the impacts
on the microbial community. Different studies showed that the biomass of microorgan-
isms in the soil can increase between 5% and 116% after addition of compost (Fuchs
2010; Tejada and Gonzalez 2006a). In mid-term applications, compost showed to
increase soil microbial biomass between 10% and 225%, whereas smaller increases in
this parameter were observed in the long-term (Table 7). Compost application also
influences the activity of soil microorganisms (Diacono and Montemurro 2010), which
are essential for the maintenance of soil functionality and fertility, as they are key play-
ers in several processes such as decomposition of organic matter and nutrient cycling.
Short-term experiments showed increases in microbial activity of 0–57%, while mid-
term trials showed that microbial activity can increase up to 263%. Microbial activity
showed variations between 1% and 7% in long-term experiments (Table 7).
Several authors signalled that compost can also influence microbial diversity.
Data determined through molecular techniques analyzing microbial DNA profiles
such as denaturing gradient gel electrophoresis (PCR-DGGE) or terminal restric-
tion fragment length polymorphism (TRFLP), showed that the effects on soil micro-
bial diversity are highly variable, ranging from reductions (−2.2%) to increases
(1.8%) (Table 7). Compost addition also alters soil microbial functional diversity as
evidenced by community level physiological profiles. Higher functional diversity
indicates that the microbial community is able to carry out a broader range of meta-
bolic reactions (Garland and Mills 1991). Short-term experiments showed that func-
tional diversity increased between 5.21% and 9.38%, and between 4.09% and 7.02%
in long-term experiments (Table 7). No data were found for mid-term experiments.
Literature covering aboveground species is limited, and results are in some cases
inconclusive. For instance, Tejada et al. (2009a) showed that compost can increase
the biomass of natural vegetation by 68.7% in degraded soils, while Bastida et al.
(2008) observed that composted sewage sludge increased natural plant biomass but
decreased significantly species diversity as compared to a non-amended control. In
contrast, Sutton-Grier et al. (2009) observed that compost addition to a degraded
wetland had limited early effects on plant communities.
3.9 Crop Nutritional Quality
Phytochemicals or secondary metabolites have recently attracted the attention of the

industry, scientists and consumers, because of their potential health benefits, Fig. 5
(Sun and Tanumihardjo 2007). Phytochemicals are produced by plants in response
Table 7 Literature review of the effects of compost on soil microbial community. Data on compost impacts on soil biodiversity is scarce and restricted to
microorganisms. Compost may decrease microbial diversity by 2% or increase it by 4%. Clear impacts are also observed on microbial biomass (3.2–242%
increase after compost application) and microbial activity (43–344% increase)
Change in
Dosage ww the diversity
Study Compost feedstock (t ha−1 y−1) Crop Type of soil Length (yr) indicator (%)
Microbial Biomass Carbon
Albiach et al. (2000) MSW 24 No crop Xerorthent sandy 5 187.5
silty loam
García-Gil et al. (2000) MSW 20 Barley Typic haploxeralf 9 10.2
MSW 80 Barley Typic haploxeralf 9 46.8
Ros et al. (2003) MSW 300 No crop Xeric torriorthents 2 225
Ros et al. (2006a; MSW 22.5 Corn-wheat-barley Loamy silt 12 8.2
2006b) Green waste 16.9 Corn-wheat-barley Loamy silt 12 10
Cattle manure 20.8 Corn-wheat-barley Loamy silt 12 3.2
Sewage sludge 12.2 Corn-wheat-barley Loamy silt 12 4.7
Tejada and González Crushed cotton gin 10 Rice Aquic xerofluvent 1 21.7
(2006a) Crushed cotton gin 15 Rice Aquic xerofluvent 1 60.9
Crushed cotton gin 20 Rice Aquic xerofluvent 1 116.3
Fließbach et al. (2007) Cattle manure 11.2 Typic hapludalf 21 38.3
Bastida et al. (2008) Sewage sludge 120 No crop Haplic calcisol 1 106.4
Tejada et al. (2009a) Beet vinasse + green- 7.91 No crop Xerollic calciorthid 3 183.3
waste vermicompost
Beet vinasse + green- 15.82 No crop Xerollic calciorthid 3 241.7
waste vermicompost
(continued)
291
292
Table 7 (continued)
Change in
Microbial Diversity H’ (PCR-DGGE)
Ros et al. (2006a; MSW 22.5 Corn-wheat-barley Loamy silt 12 −2.24
2006b)
Green waste 16.9 Corn-wheat-barley Loamy silt 12 3.73
Ge et al. (2010) MSW 25 Hapludalf silty clay nm 1.8
loam
MSW 75 Hapludalf silty clay nm 0
loam
Bacterial Functional Diversity H’ (CLPP)
Gómez et al. (2006) Household solid waste 20.74a No crop Vertic argiudoll 0.5 5.21
Household solid waste 41.49a No crop Vertic argiudoll 0.5 9.38
Ros et al. (2006a; MSW 22.5 Corn-wheat-barley Loamy silt 12 7.02
2006b) Green waste 16.9 Corn-wheat-barley Loamy silt 12 5.26
Change in
Microbial Activity (basal respiration)
Ros et al. (2003) MSW 300 No crop Xeric torriorthents 2 263.6
Pérez-Piqueres et al. Green waste compost 20% (vol) Clayey soil 10 days 0
(2006)b Spent mushroom (UK) 20% (vol) Clayey soil 10 days 344.4
Spent mushroom (Fr) 20% (vol) Clayey soil 10 days 555.6
Green waste compost 20% (vol) Sandy silty clay 10 days 57.1
soil
Spent mushroom (UK) 20% (vol) Sandy silty clay 10 days 123.8
soil
Spent mushroom (Fr) 20% (vol) Sandy silty clay 10 days 120.6
soil
Ros et al. (2006a; MSW 22.5 Corn-wheat-barley Loamy silt 12 1
2006b)
Green waste 16.9 Corn-wheat-barley Loamy silt 12 7
Cattle manure 20.8 Corn-wheat-barley Loamy silt 12 0
Sewage sludge 12.2 Corn-wheat-barley Loamy silt 12 43
Fließbach et al. (2007) Cattle manure 11.2 Typic hapludalf 21 23.8
Bas tida et al. (2008) Sewage sludge 120 No crop Haplic calcisol 1 121.7
MSW municipal solid waste, vol volume, nm not mentioned
a
Only dry weight is available in the study. We considered 35% moisture content for the ww calculation
b
Laboratory incubation
293
Fig. 5 Bioflavonoid, a bio-active compound of food, is presented as a health value in a Catalan

grocery. Translation of the poster: “Bioflavonoid is a phytochemical component that is only avail-
able in raw fruits and vegetables. It is a true natural medicine, highly used by the pharmaceutical
industry, as it is very important to fight cell aging” (Photo by Joan Rieradevall)
to biotic or abiotic sources of stress. The type and amount of metabolites produced
is influenced by genetic and agronomical factors (Gratacós-Cubarsí et al. 2010). In
largely fertilized agricultural fields, plants have typically a quick growth, thereby
developing low concentrations of antioxidants and some vitamins as a consequence
of this “dilution effect” (Benbrook 2005).
From a crop management perspective, studies suggested that organic farming has
a tendency towards nutritional superiority but lower yields compared to conventional
agricultural (Benbrook 2005, 2008; Lairon 2010). Based on 236 case studies, Benbrook
et al. (2008) reported that the organic agricultural products were nutritionally superior
in 61% of the cases, while conventional products contained more nutrients in 37% of
the cases. De Pascale et al. (2006) demonstrated that both the farming system and N
rate affect carotenoid content and antioxidant activity of tomato fruits.
In this context, few studies showed the beneficial effects of compost with respect
to the level of phytochemicals in crops (Table 8). Higher levels of several antioxi-
dants when compost was used in pot experiments with dosages above 50% in vol-
ume were reported by Wang and Lin (2003). Coria-Cayupán et al. (2009) assessed
a higher content (between 20% and 35%) of chlorophylls and carotenoids in lettuce
grown with urban or fruit-and-vegetable waste composts, but lower contents or non-
significant differences for phenol compounds and antiradical activity. Martínez-
Blanco et al. (2011) also found up to 76% and 24% increase in the content of sinapic
acids and phenols, respectively, in cauliflowers when compost was applied to soil.
Table 8 Literature review of crop nutritional quality after compost application in soils. The effects are very variable and depend on specific phytochemical and
local conditions
Increase in the
nutritional content
from options without
Study Compost feedstock Dosage ww Type of soil Crop Length (yr) Phytochemical compost (%)
Wang and Dairy and poultry 50% soil + 50% – Allstar and 1 Ascorbic acid 8.9–15.03
Lin (2003)a manure + livestock compost honeoye Dehydroascorbic acid ns
bedding + landscape strawberry Reduced glutathione 4.7–12.7
trimmings + green- Oxidized glutathione ns
house dis- Ellagic acid 16.0–140.0
cards + wood
p-coumaroylglucose 23.9–65.8
chips + leaves
Dihydroflavonol 44.6–116.0
Kaempferol 12.5–32.6
3-glucoside
Kaempferol 22.1–51.0
3-glucuronide
Cyanidin 3-glucoside 39.5–76.6
Pelargonidin 17.2–25.9
3-glucoside
Cyanidin 3-glucoside- 31.0–71.8
succinate
Pelargonidin 14.3–28.5
3-glucoside-
Succinate
Coria-Cayupán biowaste + MSW 10 t ha−1 – letucce 0.15 Chlorophylla 15.3–21.7
et al. (2009) Chlorophyllb
15.6–16.4
b-carotene 6.7–20.0
(continued)
295
Table 8 (continued)
296
Increase in the
nutritional content
Lactucaxanthin ns
Lutein 7.7–23.1
Violaxanthin 11.5–26.9
Neoxanthin 40.0–170.0
VitA 7.3–23.6
Caffeic acid ns
Coumari cacid ns
Martínez- biowaste 200/100 t ha−1 b typic cauliflower 1c Sinapic acid ns/ns
Blanco et al. Xerothent 1,2-disinapoyl-
(2011) diglucoside
74.1/ns
1-sinapoyl-2- ns/ns
feruloyldiglucoside
1,2,2¢-trisinapoyldig- 88.8/ns
lucoside
1,2¢-disinapoyl-2- 76.5/ns
feruloyldiglucoside
Totalphenols 24.1/ns
Glucoiberin ns/ns
Sinigrin ns/ns
Glucoraphanin
ns/ns
Progoitrin
ns/ns
Glucoalyssin
−45.5/-45.5
Increase in the
nutritional content
Glucoiberverin ns/ns
Glucoerucin 150.0/ns
4-OH-Glucobrassicin ns/ns
Glucobrassicin
ns/ns
Metoxiglucobrassicin
ns/ns
Neoglucobrassicin
ns/ns
Kaempferol-3- 143.3/ns
diglucoside-7-
diglucoside
Quercetin-3- ns/ns
diglucoside-7-
glucoside
Kaempferol-3- ns/ns
diglucoside-7-
glucoside
ns no significant differences
a
Pot experiment
b
Two options were considered. The results in the last column are separately presented for the two dosages
c
The crop lasted 112 days. The crop is the fourth crop in a cultivation rotation. The crop was cultivated 313 days after compost application
297
Accordingly, increasing nutrient contents are often observed in organic production

and when compost is employed. However, general figures cannot be drawn, as the
response varies depending on the compounds being assessed, the crop type, and the
environment conditions (mainly soil and climate), while compost affects more than
one compound at the same time.
3.10 Summary of the Benefits of Compost Application
According to all the literature summarized in the Tables 1, 2, 3, 4, 5, 6, 7, and 8 and

explained in the previous sections, an overview is provided in Table 9 for each of the
benefits.
Effects in the mid- and long-term were only reported for nutrient supply, carbon
sequestration, crop yield, soil erosion, soil biodiversity and soil workability; whereas
for the other four potential benefits only short-term data could be reported.
For three out of the nine benefits assessed – pest and disease suppression, biodiver-
sity increase and nutritional content increase –, although they were proved, it was not
feasible to summarize the effect in a unique data range. In these cases compost is
affecting more than one indicator at the same time and the intensity of the effect is
different for each one – e.g. compost application is inducing and increase of a large
number of nutritional compounds while non-significant effects are being detected for
other compounds.
Significant effects due to compost application were averaged for all the potential
benefits, except for weed suppression. In addition, for three of the assessed effects
the share of studies with non-significant results was relevant: for crop yield, more
than 60% of the case studies did not detect differences when compost was applied;
non-significant effects were detected for soil moisture content for low rates of com-
post; and finally, crop nutritional quality was not relevantly different for a third of
the case studies included. These were also the benefits with higher disparity in the
measured effects among the results.
Some of the possible reasons for such variation are the composition of the initial
waste used in compost production, the management of the composting process,
compost maturity, crop management, and the different analytical techniques used to
evaluate the same parameters. The dosages of compost applied as well as the exis-
tence of a lag of time between compost application and crop sowing, are also key to
explain the strength of the observed effects.
4 Quantification and Impact Assessment
In the following sections, benefits from compost application are quantitatively

described within an LCA perspective. Firstly, the consequential modeling for each
benefit is briefly described, followed by quantification of the substituted or saved
Table 9 Summary of the potential benefits of compost application in the short-, mid- and long-term retrieved from the literature review
Short-term mid-term long-term
(<1 year) (<10 year) (<100) Number of case
Related section Effect Unit Min. Max. Min. Max. Min. Max. studies reviewed
2.1 N mineralized % of N applied 5 22 40 50 20 60 5 RW ,1 CS
P mineralized % of P applied 35 38 90 100 90 100
K mineralized % of K applied 75 80 100 100 100 100
2.2 C sequestered in soil % of C applied 40 53 30 2 16 14 CS
2.3 Weed suppression – ns ns – – – – 1 RW
Pest and disease – nad nad – – – – 1 RW
suppression
2.4 Crop yield gain % from min. Fert. −495 0 −71 52 – – 77 CS
2.5 Soil erosion decrease % soil loss – – −5 −36 – – 4 CS
D structural or aggregate 29 41 0 63 – – 8 CS
stability (%)
2.6 Soil moisture increase WHC increase (%) 0 50 – – – – 21 CS
PAW increase (%) 0 34 – – – – 10 CS
2.7 Improved workability D soil bulk density (%) −2.5 −21 −0.7 −23. −20 21 CS
2.8 Biodiversity increase D Microbial diversity (%) – – – – −2 4 2 CS

D Microbial biomass (%) 22 116 10 242 3.2 100 8 CS
D Microbial activity (%) 0 344 – 264 0 43 5 CS
2.9 Nutritional content – nad nad – – – – 42 CS
increase
WHC water holding capacity, PAW plant available water, ns no significant differences, nad no average data because of complexity of available dataset, “–”, no
reported effects, RW review, CS case study
Negative values indicate a decrease in the indicator
299
COMPOST BENEFITS Impact categories currently

accepted
Nutrient supply
Acidification
Carbon sequestration Aquatic eutrophication
Weed, pest and desease Global warming

suppression
Ecotoxicity
Crop yield*
Impact categories under- Energy consumption
development
Soil erosion Ozone depletion
P resource
Soil moisture content Human toxicity
Biodiversity
Soil workability Ozone formation

Land use and soil quality
Resources consumption
Water use and quality Soil biological properties
and biodiversity
Terrestrial eutrophication
Crop nutritional quality*
Fig. 6 Midpoint life cycle assessment (LCA) impact categories involved in the evaluation of the
potential benefits of compost use-on-land. *It has to be included in the LCA using the adequate
functional unit
process based on up-to-date knowledge and data (Table 9). Finally we address the
impact categories in existing impact assessment methodologies which are most
affected when considering compost application (Fig. 6). Other methodological
issues are addressed in the Discussion section.
4.1 Nutrient Supply
In an LCA context, the supply of nutrients with compost substitutes the use of
mineral fertilizers, whose industrial production and transport is thus avoided. The
amount of substituted fertilizers depends on the content of nutrients of the compost
and their utilization rate. The substitution is often regulated by legislation (Hansen
et al. 2006). Boldrin et al. (2009) reported that final utilization efficiencies for N, P
and K are in the order of 20–60% for N, 90–100% for P, and 100% for K. Using the
nutrient contents presented in Sect. 3.1, the potential amount of inorganic fertilizers
replaced may be within the range of 1–13 kg of N, 1–5 kg of P, and 5–14 kg of K
per ton of compost applied.
A life cycle inventory for fertilizer production includes the use of materials and
energy, and emissions to different compartments, which would typically result in
potential impacts on Resource Depletion, Global Warming, Human- and Ecotoxicity,
and Eutrophication categories. Datasets for N-P-K fertilizers are reported by differ-
ent sources, as for example Wind and Wallender (1997), Hansen et al. (2006) based
on Patyk and Reinhardt (1997) and Audsley et al. (2003), and Ecoinvent based
mainly on Davis and Haglund (1999).
Furthermore, compost is considered as an effective option for phosphorous recy-
cling (Cordell et al. 2009), which is a growing issue as a consequence of the fore-
seen shortage of mineral P for agriculture fertilization (Syers et al. 2008). The use
of a renewable P source rather than inorganic non-renewable supply is of great
importance and this reduced raw resource consumption might be quantified during
impact quantification.
4.2 Carbon Sequestration
Sequestration of C into soil can be seen as removal of C from atmosphere. Thus,

once the amount of C sequestered is estimated, the value can be translated into
saved CO2 emissions, using a conversion factor of 44/12, based on molar relation,
which can be entered in the inventory. As previously mentioned, the time-horizon
used in the assessment plays a crucial role when estimating the benefit from carbon
sequestration. A time frame of 100 years is considered to be relevant for estimating
contributions to global warming (Favoino and Hogg 2008). Boldrin et al. (2009)
reported that the benefits from C retained in soil 100 years after the addition of biowaste
compost is between 2 and 79 kg CO2-eq. t−1. Higher values, 279 kg CO2-eq. t−1, were
reported by ICF (2005). Most likely, this large variability is due to the synergistic
effect of the different abovementioned environmental and site-specific factors,
meaning that estimations should be done on a case-to-case basis.
4.3 Weed, Pest, and Disease Suppression
When the application of compost reduces the incidence of weeds, pest and diseases, the
use of herbicides or pesticides can be reduced or avoided. The avoided use can conse-
quently be credited to the system as an environmental saving. The studies reviewed
indicate that the use of compost might increase, in most cases, the health of crops through
resistance towards certain diseases, especially fungal diseases. However, these benefits
are so case-specific that it is not possible to provide any general figures, regarding both
the amount and the type of pesticides saved. For example, ROU (2007) estimated that
2–3 L per ha of 360 g/L glyphosate could be saved when using garden waste compost as
mulch, however no benefits were estimated if compost was used as soil amendment.
When pesticides are saved, environmental benefits are related to both the avoided
production/transportation and to the avoided use of the pesticides. Inventory data
covering transport and production of pesticides can be found in different databases
such as Ecoinvent, PestLCI and GEMIS. Assessment of the environmental effects
induced from pesticide utilization requires the use of exposure-fate-toxicological
models. In fact, on one hand pesticides are purposefully emitted to the environment
to control soil and air-borne plant pests, and diseases, on the other hand they can
also reach non-targeted life forms via wind drift, evaporation, leaching, and surface
run-off, with consequent potential toxic effects on human or environment (Brentrup
2004; De Bertoldi 2010; Juraske et al. 2009).
Unfortunately, dynamic and realistic models capable of predicting chemical fate
taking into account all the factors are scarce (Birkved and Hauschild 2006) and
require exhaustive information of the pesticide’s characteristics and application
conditions, which in many cases are unknown (Antón et al. 2004). An available
model is PestLCI, developed for inventorying pesticide emissions to the different
environmental compartments, and based on information which will normally be
available to the model user (Birkved and Hauschild 2006). Potential environment
impacts from production/transportation and use of pesticides can be assessed using
existing impact categories. The most relevant impact categories related to pesticides
are the Toxicity categories, both Human- and Ecotoxicity.
4.4 Crop Yield
Increased yield as a consequence of compost application results in avoided agricul-

tural production, and thus the involved burdens. The increase should be measured
relative to the yield otherwise obtained by using the amount of fertilizer already
substituted, in order to do not count twice the effect of the compost application.
From a consequential LCA point of view, this can have different consequences at a
system level, depending on existing agricultural constraints. If arable land in a cer-
tain area is not constrained, the benefit is linked to avoided use of material and
energy needed for the crop production. In the most likely regime of constrained
arable land, the increased yield would have an effect on both intensification and
expansion of agricultural production, and ultimately will prevent indirect land use
changes, which are for instance a major source of greenhouse gas emissions through-
out the life cycle of biofuels (Thamsiriroj and Murphy 2010).
The LCA modelling of increased or decreased agricultural productions is typi-
cally case-specific. Market driven mechanisms need to be taken into consideration
when identifying both the specific crop directly affected by the compost-induced
increased yield and the indirect land use changes. For both effects, production
inventories can then be used to credit the system for the avoided productions.
Depending on the specific area and crop, most of the impact categories are influenced
when agricultural production is involved.
4.5 Soil Erosion
As mentioned earlier, the application of compost could prevent soil erosion and
thereby avoid losses of arable land. Probably due to the absence of data, the reduction
in soil erosion was not taken into account in previous LCA of compost use-on-land.
As shown in Table 1, losses of soil could decrease between 4% and 66% with the
application of compost, depending on the time horizon considered. A more precise
quantification is possible for specific conditions taking into account for instance cli-
mate, application rate, and type of soil.
Two approaches are possible during inventory modelling. One option is to model
the avoided losses within traditional LCA impact categories. Here the consequential
modelling should identify the agricultural production affected by the losses of ara-
ble land. Assuming a constrained agricultural production at a system level, the mod-
elling is then done similarly to “crop yield” section, meaning that the consequences
of intensification and/or land expansion are included in the assessment. The second
option is to consider soil as a resource, thus either including loss of soil in the inven-
tory as ‘resource depletion’ (Cowell and Clift 2000) or introducing ‘soil erosion’ as
an independent endpoint impact category (Buratti and Fantozzi 2010). This issue is
further elaborated in the discussion session.
4.6 Soil Moisture Content
Actual water necessities of the crop are not affected by compost application. The
potential benefit of compost is to raise the capacity of soil to retain green water, i.e.
rainfall and irrigation water stored in the soil as soil moisture, in order to reduce
irrigation and consumption of blue water, i.e. water from surface and groundwater
resources. This may result in two distinguished consequences: on one hand, blue
water is saved; on the other hand, because more green water is available, crop yield
could increase in those areas where irrigation water is not available. However,
there is not a direct relation between the retention capacity and the amount of water
saved, depending on site conditions, water demands of the plants, management
practices, and previous moisture content of soils. The theoretical saving of irriga-
tion water could be calculated for a particular case study, if all these data are avail-
able, as done for example in ROU (2007). Alternatively, results from experiment
trials could be used. The only study on this issue was carried out by Ngoundo et al.
(2007), who reported that, in both tropical humid and semi-arid regions, compost
application was a reliable way of saving water. Authors concluded that compost
application on water-intensive crops increased the net irrigation depth, thereby
reducing the need for irrigation.
Environmental burdens from irrigation water supply are linked to water extrac-
tion, transport, and distribution in the field, including electricity, pumps, pipes, etc.,
and are found in several inventories, also at a regional level. Potential impacts from
these processes are typically those related to energy supply and consumption, thus
Global Warming, Acidification, and Eutrophication. There is indeed a growing con-
sensus on the fact that water should be considered a resource, and thus its consump-
tion included in LCA as Resource consumption (or depletion) (Berger and Finkbeiner
2010; Heuvelmans et al. 2004; Núñez et al. In press 2012a). This issue is further
elaborated in the discussion section.
4.7 Soil Workability
Improved soil workability can potentially decrease energy requirements for agricultural
operations (Favoino and Hogg 2008; ROU 2007). Soil ploughing typically involves
large consumption of energy and fuel, being one of the farm operations with most
environmental impact (Lal 2004b). Hillier et al. (2009) in a survey carried out in 57
farms in Scotland (UK), estimated that the carbon footprint of ploughing operations
was on average 15.2 kgCO2-eq. ha−1 (corresponding to approx. 5.7 Ldiesel ha−1). Lal
(2004b) carried out an extensive literature review and reported that estimates of
emissions for different tillage operations were between 2 and 20 kg CO2-eq. ha−1
(i.e. 0.75–7.5 Ldiesel ha−1).
In an 8-year long study, it was reported that, under a 100 t ha−1 application of
stockpiled and rotted manure on a corn field, the plough draft was reduced 27–38%,
resulting in 13–18% reduction of fuel consumption (McLaughlin et al. 2002). The
fuel consumption accompanying the organic amendments was attributed to an
improvement in soil tilt and quality related to the increase in SOM. No other studies
were found linking compost application and fuel consumption for agricultural oper-
ations, meaning that more comprehensive data are needed to be able to relate, for
example, fuel consumption with soil bulk density.
Reduced fuel consumptions can be credited to the system as avoided use of die-
sel. Inventory data for diesel consumption, including provision, are available in sev-
eral databases and published report. Avoided diesel consumption mainly affect
Global Warming impact category, while avoided emissions of nitrogen oxide could
also have an influence on Acidification and Eutrophication.
4.8 Soil Biological Properties and Biodiversity
Changes in soil biodiversity after compost addition might influence either positively
or negatively the services delivered by the ecosystem, with consequences in terms
of impacts associated to the substitution or compensation of those ecosystem ser-
vices. Soil organisms are the basis of several ecosystem services and regulate soil
processes such as, for instance, soil hydrological processes, nutrient cycling, and
the incidence of pests, parasites and diseases (Brussaard et al. 2007).
Alterations in the system service in connection to biodiversity changes could be
modelled within the traditional categories if those changes could be quantified in the
inventory. If, for example, increased biodiversity could be directly related to better
nutrient cycling and lower need for fertilization, then the benefits from increased
biodiversity could be modelled in terms of reduced production of fertilizers.
However, data linking compost use, biodiversity and ecosystem services are non-
existing – apart from a first attempt of establishing a preliminary relation by
Nemecek et al. (2011) –, meaning that such quantification is currently not feasible.
In addition, general figures could not be established in all case, as the effects of land
management practices are highly variable depending on regional and scale dependent
factors (Bengtsson et al. 2005). An alternative approach is to consider biodiversity

and ecosystem services as independent endpoint categories when assessing the
environmental impacts of land management alternatives (Zhang et al. 2010). Some
recent initiatives have established baseline diversity indices for different soil organ-
isms and under different soil uses that can be used as a reference to evaluate the
impacts of compost on soil biodiversity (Cluzeau et al. 2012).
4.9 Crop Nutritional Quality
The differences in the nutrient content of crops do not directly affect resources con-
sumption or emissions per hectar or ton yielded. Nevertheless, different nutrient
contents in products can have a repercussion on the LCA modelling depending on
how the functional unit is defined. If the functional unit is based on yield, i.e. mass,
volume, surface, no consequences on resource consumption or emissions need to be
modelled in the inventory. However, when the functional unit includes qualitative
aspects such as nutritional and/or economic value, increased nutritional level of a
food product may have as a consequence that lower amounts are needed. In general
terms, including qualitative aspects in the functional unit, would have an effect on
the agricultural production, which could be modelled similarly to what is described
under the “crop yield” section (Marshall 2001; Martínez-Blanco et al. 2011).
5 Discussion
Regarding the proof of the effects and the environmental assessment – including
quantification and characterization – of the benefits of compost application to soils,
four different scenarios were identified:
• Scenario 1: The positive effects of compost application are proved, effects are
quantifiable, and tools for their consideration with LCA are available. This
includes nutrient supply and carbon sequestration which are, and should be,
included in LCA studies.
• Scenario 2: The benefits are proved, but their magnitude is too variable as a con-
sequence of the synergetic effect of many factors. Thus, inventory data cannot be
unambiguously quantified. Impact categories and characterization factors exist
for most of the benefits.
• Scenario 3: The benefits are proved and quantifiable. However, corresponding
characterization factors and/or impact categories are non-existing.
• Scenario 4: Benefits are not fully proved and thus their inclusion in the modeling
is not yet feasible.
Therefore, from all the nine benefits proposed, only two are proved and the
quantification and assessment are possible, while different research efforts are
required in the rest of the benefits for a full assessment. In the following sections,
open issues related to quantification and characterization are discussed.
5.1 Quantification: Improved Modelling
Review of the existing literature, shows that in the future, modeling of effects from
compost application should be improved in some areas in order to adequately assess
its benefits.
First, LCA models are typically linear steady-state models of physical flows
(Guinée et al. 2002). Fluxes of nutrients and pollutants after compost application to
soil are not linear in most of the cases. This also applies, for instance, to repeated
applications of compost: LCA studies typically look at the effects of a single appli-
cation over 100 years, while the cumulative effects on several applications may not
be linear with the amount of compost added.
Second, LCA models assume that impacts depend on the compost characteristics
while they rarely include environmental parameters as determining factors. This
links with the necessity of coupling LCA and agronomic models to gain a more
precise picture.
Third, the amount of plant nutrients contained in compost is normally mod-
elled for as a benefit. However, the use of compost could in some cases result in
excessive application of P and K with respect to N, which is usually used for com-
post dosage calculation. The subsequent release of this unbalanced amount of
nutrients could result in impacts to the environment and should be thus included
in the LCA modelling through a more thorough mass balancing of the nutrients.
To avoid the negative environmental impacts and the resources losses, compost
doses should be better calculated based on crop P and K necessities and to add
nitrogen in other forms.
Fourth, for LCA of the agricultural sector, the functional unit is typically defined
per area used or product yield. The purpose of the functional unit is to provide a
reference to which the inputs and outputs are related, and to ensure comparability of
LCA results (ISO 2006). As different functional units can lead to different results
for the same product system (Deytieux et al. 2012; Marshall 2001; Martínez-Blanco
et al. 2011), there may be a need for a qualitatively more precise definition when
dealing with compost application, especially in those cases where the product qual-
ity is affected. Better definitions could, for example, include the economic value or
the nutritional content of a product (Hayashi et al. 2006; Mourad et al. 2007; Reap
et al. 2008; Schau and Fet 2008). A more accurate definition dealing with nutritional
differences may include a combination of nutritional quality and quantity or yield.
This was for example done by Charles et al. (1998) and Audsley et al. (2003), where
the functional unit was defined as “1 equivalent ton grain with 12–13% protein”.
This involved the use of marginal productions to adjust the overall output of the
system under assessment. However, identifying reference or recommended nutrient
content for most crops may be a controversial issue.
Finally, the choice of the time horizon of the LCA should be harmonized. The
studies reviewed showed in fact that such choice is in many cases very important, as
both the foreground and background effects of compost application vary largely
depending on the time frame.
5.2 Characterization: Additional Impact Categories

and Proposed Modifications
As highlighted in the text, when applying LCA for assessing use-on-land of com-
post, available impact assessment methodologies may not properly deal with some
issues, and thus new impact categories or modifications of the current ones may be
needed in the future to allow for a more holistic assessment. The proposals should
deal with depletion of P resources, biodiversity, loss of arable soil, and consumption
of water.
Depletion of P as a resource is currently modelled similarly to other natural
resources. However, most metals, chemicals, even petroleum, could be replaced by
other resources in much of their functions, while this is not the case of phosphate.
A revision of the characterization factors is thus needed for the assessment of non-
replaceable non-renewable resources such as P. In this respect, the ReCiPe model
adds special value to resources. It is based on the geological distribution of mineral
and fossil resources, and assesses how the use of these resources causes marginal
changes in the efforts to extract future resources (Goedkoop and Spriensma 2000).
Assessment of the impacts of land management on biodiversity can be based on
particularly endangered species (Nemecek et al. 2011). Other authors estimated the
changes in biodiversity through the assessment of one relevant indicator group. For
example, Weidema and Lindeijer (2001) developed an indicator that includes species
richness of vascular plants, inherent ecosystem scarcity, and ecosystem vulnerability.
Similarly, the Eco-Indicator 99 measures ecosystem quality based on the potential
disappeared fraction of vascular plants (Goedkoop and Spriensma 2000). Other
methods estimated the change in the overall number of species per year (Suer and
Andersson-Sköld 2011) or the change in the number of species in the affected area
as compared to the regional level (Köllner 2002). Other authors have included biodi-
versity and ecosystem services as midpoint categories within the endpoint category
Land Use (Milà i Canals et al. 2007; Udo de Haes 2006). As different options exist,
a harmonization may be needed in order to develop a consensus methodology.
Soil loss involves the loss of cultivable land but also the loss of SOC, plant nutri-
ents, as well as the associated plant, animal and microbial biodiversity (Cowell and
Clift 2000). Loss of soil can thus be included in some of the abovementioned impact
categories. However, for a more comprehensive assessment the loss of soil mass
could be considered as the loss of a resource and included in the inventory as
Resource Depletion (Cowell and Clift 2000; Núñez et al. In press 2012b). The amount
of soil loss could, for example, be first normalized against the available soil reserves
(Cowell and Clift 2000; Núñez et al. In press 2012b) or against the estimated soil
eroded in the region of study within a specific time frame (Buratti and Fantozzi
2010). Further weighting can be based on the exergy use for the backup technology
needed to maintain the soil productivity (Núñez et al. In press 2012b), or by relating
the amount of soil loss to reference values of soil erosion where no significant decline
in soil productivity is observed (Buratti and Fantozzi 2010). In alternative, soil ero-
sion can be included within the impact category Land Use, whose characterization
factors are based on soil quality indicators such as SOM, structure, heavy metals,
biodiversity, aesthetic value, etc. (Brentrup 2004; Mattsson et al. 2000).
Depletion of water resources is gradually gaining importance, particularly in cer-
tain geographical regions. There is currently only a preliminary scientific consensus
about the parameters to consider and the methodology to follow (Núñez et al. In
press 2012). Methodological issues concerning impact assessment methods include
the types of water use accounted for, the inclusion of local water scarcity conditions,
and the differentiation between watercourses and quality aspects (Berger and
Finkbeiner 2010). While available inventories account for the total volume of water
used within the life cycle of products (Hoekstra et al. 2009), at the impact assess-
ment level, more emphasis is given to the blue water consumption, i.e., consump-
tion from surface and groundwater resources. However, from an environmental
point of view interest of green water consumption by crops is also important because
of its influence in ecosystems (Berger and Finkbeiner 2010).
6 Conclusion
Use of biowaste compost on land can have beneficial effects on the plant-soil sys-
tem. Most of these benefits have been so far excluded from LCA studies, mainly
because of scarcity of data or lack of appropriate impact assessment methods. In
this study, a literature search was carried out in which nine benefits of compost
application were identified and studied its influence and consequences on potential
environmental problems. Availability and quality of the data for quantification dif-
fered largely among the assessed benefits, with no data or large variability in the
observed benefits. Data concerning long-term effects of compost, which are relevant
for LCA purposes, were particularly scarce. Therefore there is a need for more long-
term studies or estimations.
When data were available, local conditions and ecosystem complexity were the
main obstacles for a precise quantification. Furthermore, most of the studies consulted
used different compost doses – annually or a unique application – and for different
periods of time, thereby increasing the difficulty of quantifying the benefits.
A discussion on the suitability of currently available impact assessment method-
ologies indicated that additional impact categories may be needed for a comprehen-
sive assessment of compost application. For two of the nine benefits – nutrient
supply and carbon sequestration – the review showed that both quantification and
impact assessment of the effects could be performed, meaning that these two benefits
should be regularly included in LCA studies. For four of the nine benefits – “increase
in crop yield” in crop yield, soil workability, “crop nutritional change”, and
“enhancement of soil biological properties and biodiversity” –, quantitative figures
could not be provided, either because of complete lack of data or because the effects
are both very variable and too depending on specific local conditions. For “soil ero-
sion” and “soil water content” effects could be quantitatively addressed, but avail-
able impact assessment methodologies were considered unsuitable to
comprehensively evaluate the implication of compost application with regards to
these two benefits. Finally, based on the available literature, “suppressive effects of
compost on weed, pests, and diseases” could not be generally proved.
Acknowledgements The authors would like to thank the Spanish Ministerio de Educación for the
research scholarship (AP2008-02954) awarded to Julia Martínez Blanco and the financial support
by an «Ángeles Alvariño» fellowship from Xunta de Galicia to Cristina Lazcano.
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es900548a
Biocontrol of Plant Pathogens Using Plant
Growth Promoting Bacteria
Pratibha Prashar, Neera Kapoor, and Sarita Sachdeva
Abstract Food is the primary requirement for the survival of mankind. Therefore
food has always been an issue since prehistoric times. Food production through
agriculture is a key process to meet the world food demand. In the last four to five
decades global agricultural production has been drastically improved by introduc-
tion of improved crop varieties and development of modern agricultural tools.
However, more than 25% of the crop yield is lost every year due to various kinds of
plant diseases. Most critical plant diseases are caused by soilborne plant pathogens
such as fungi, virus and nematodes. Effective measures are thus highly needed to
avoid this crop loss. So far, the use of chemical pesticides has been the method of
choice to tackle this problem. However, the unwarranted use of chemical tools to
enhance the crop yield and control plant diseases has resulted in irreversible loss of
soil quality along with serious health and environmental problems. Moreover, alter-
native controls are needed due to growing public concern about toxic agrochemicals
and stringent laws. Microbes are a reasonable option to develop ecofriendly agricul-
tural tools to replace chemical pesticides.
Here we review the role of a special class of soil bacteria, called plant growth
promoting rhizobacteria (PGPR). PGPR live in the rhizospheric sites in soil, i.e. in
immediate vicinity of the plant roots, and exerts several beneficial effects on the
plants, directly or indirectly. PGPR have inherent antagonistic properties against
P. Prashar (*)
Department of Biotechnology, FET ,MRIU, Faridabad India
School of Sciences, IGNOU, New Delhi, India
e-mail: pprashar@gmail.com
S. Sachdeva
Department of Biotechnology, FET, MRIU, Faridabad, India
e-mail: sarita.fet@mriu.edu.in
N. Kapoor
School of Sciences IGNOU, New Delhi, India
e-mail: neerakapoor@ignou.ac.in
320 P. Prashar et al.
soilborne plant pathogens under natural conditions. The major points discussed here
are: (1) biocontrol of plant pathogens as an alternative to chemical control methods.
(2) The current market status of various biocontrol products with a noticeable
increase of 80% market share in 5 years. (3) The success of PGPR to protect many
plant species, against diverse range of plant pathogens. (4) Mechanisms underlying
the control and inhibition of soilborne plant pathogens including antagonistic
activities such as production of antibiotics, offering stiff competition to the patho-
gen for nutrients and niches in the rhizosphere, parasitism of the pathogen and
induction of systemic resistance in the plants against diseases.
Keywords Sustainable agriculture • PGPR • Biocontrol • Soil borne plant

pathogens
1 Introduction
Attaining food security through sustainable approaches is a blazing issue, world

over. Though dramatic improvements have been made in terms of crop production
in last few decades, plant diseases have remained a major hindrance in achieving
the targets. Soil borne phytopathogens are well established as the cause for many
devastating plant diseases leading to 25–100% crop loss every year, across the
globe (Glick and Bashan 1997). These are defined as the pathogens that cause
plant diseases via inoculums that come to the plant through soil (Koike et al.
2003) and they may complete their entire life cycle within the soil, or may spend
part of it on the phyllosphere. Soil borne phytopathogens may include a diverse
group of organisms including bacteria, fungi, virus and nematodes; however fun-
gus are the most critical amongst them as they are known to cause large number
of soil borne diseases many of which lead to most serious agricultural losses
annually (Agrios 2004; Koike et al. 2003). Most important soil-borne fungi
include Fusarium sp., Phytophthora sp., Pythium sp., Rhizoctonia solani,
Sclerotinia sp., Slerotium rolfsii, Thielaviopsis basicola and Verticillium dahlia
(Jeffery et al. 2010).
Currently used approaches for the management of soil borne phytopathogens to
reduce this loss, include cultural practices like crop rotation, breeding for resistant
plant varieties and use of chemical control agents (pesticides) (Cook 1993) and
practices like fumigation, steam treatment, and solarization of soils to some extent
(Gamliel and Katan 1992). However, these are insufficient to control root diseases
of important crop plants. Moreover, owing to the deleterious effects on human and
environment health, general public, apprehension over the undue use of chemical
agents in agriculture sector has raised the alarm to hunt for safer and effectual
options. Many microbial agents, plant growth promoting rhizobacteria (PGPR) in
particular, exhibit inherent biological mechanisms to control the pathogens. PGPR
are the bacteria which reside in the rhizosphere of plant and affect it in a favorable
manner. Rhizosphere is the thin layer of soil that surrounds the roots of plant and
is influenced by their activities. Rhizosphere bacteria enjoy a close association
Biocontrol of Plant Pathogens Using Plant Growth Promoting Bacteria 321
Depletion of soil quality
Depletion of environment health
Chemical pesticides Ill effects on human and animal health
Development of resistance among pathogens
Improvement of soil quality
No adverse effect on environment
Biocontrol agents
No ill effect on human and animal health
Development of resistance not observed
Fig. 1 Comparison of chemical pesticides with biocontrol agents. Due to their non-biodegradable
nature, chemical pesticides tend to accumulate in soil and water thereby adversely affecting their
quality. Ill effects on human and animal health result due to biomagnification process, man being
at the top of food chain is most prouncely affected
with the plant and hence are the most suitable candidates to be explored for their
potential to control soil borne plant pathogens. They have been shown to play a
pivotal role in improving the plant growth and health through mechanisms like
enhanced nutrient uptake from soil, production and release of phytohormones, ele-
vating the tolerance capacity of the plant to environmental stresses and at the same
time they adversely affect deleterious organisms (phytopathogens) in the soil
including fungus, virus and nematodes, thereby controlling the diseases. Huge vol-
ume of literature has been generated in last two decades that establishes the role of
PGPR as biocontrol agents for a variety of soil borne pathogens. Thus biocontrol
methods utilizing antagonistic microorganisms associated with the plant rhizo-
sphere offer an attractive and feasible option to develop alternative agricultural
tools for control of soil borne plant pathogens and hence replace/supplement the
chemical pesticides (Fig. 1).
2 Biocontrol
In the last five to six decades the concept of biocontrol has evolved from an area of
basic research to a commercially established and successful tool in integrated dis-
ease management programmes. Significantly large number of scientific studies have
reported the potential uses of PGPR for stimulating plant growth and control of soil
borne diseases in the last few decades.
Definition of biocontrol differs depending on the target, number; type and source
of biological agents, and the level of human intervention. Most simply, biological
control is the suppression of damaging activities of one organism by other
organism(s), usually the natural enemies.
An earliest definition of the term was given by DeBach (1964) as “the action of
parasites, predators, or pathogens in maintaining another organism’s population
density at a longer average than would occur in their absence”. However over the
years it has been modified several times (Johnson and Carl 1972; Baker and Cook
1974; Cook 1988) to encompass the variety of tools and mechanisms employed for
pathogen control as well as range of pathogen suppression. In case of plant diseases
the term applies to the use of microbial antagonists to suppress diseases as well as
the use of host specific pathogens to control weed populations (Cook 1993). PGPR
in which the antagonistic activities towards soil borne pathogens are more devel-
oped as compared to their direct plant growth promotion potential are termed as
biological control agents (BCA) or biopesticide (Haas and Défago 2005). They
exhibit their antagonistic properties against phytopathogens, generally fungus, due
to a variety of mechanisms like production and release of various antimicrobial
compounds, offering stiff competition to the pathogens and inducing systemic resis-
tance in the plants. A diverse range of microbes, including plant-associated prokary-
otes and eukaryotes possess this trait. Among the prokaryotes the most successful
antagonists of soil-borne pathogens, so far, include Agrobacterium, Bacillus,
Streptomyces, Burkholderia and Pseudomonas (Sakthivel and Gnanamanickam
1987; Montealegre et al. 2003; Barea et al. 2005; Prapagdee et al. 2008; Cawoy
et al. 2011). Hence soil microflora, PGPR in particular has been exploited in the
recent past to compete against chemical control agents for controlling plant patho-
gens of various kinds and they have proved their worth as indicated by increasing
market share of biocontrol agents, in the recent years.
3 Commercial Scenario
Presently, among the prevalent biopesticides in the market, microorganism-based

products hold a significant share of 30% of the total sales and over the years market
share of biopesticides as a whole, has grown from a meager 0.2% in 2000 to 2.5%
in 2005, which is further expected to grow 15%, annually (Cawoy et al. 2011). At
the same time, a downward trend has been noticed in case of chemical based pesti-
cides since 2000, with an expected decline rate of 1.5% per year (Thakore 2006).
Further, factors like higher prices and stringent regulations for use of chemical
pesticides, the easier registration process for biopesticides coupled with the envi-
ronmental incentives, will enhance the chances for successful commercialization of
biocontrol products (Slininger et al. 2003). Developing nations, like India and
African countries, have a great potential for the use of microbial inoculants.
Amongst the microorganism-based biopesticides, bacterial products constitutes
the major part (Shoresh et al. 2010) and Bacillus thuringiensis alone accounts for
more than 70% of total sales (Cawoy et al. 2011). Most successful commercial
products are based on Bacillus thuringiensis, Bacillus subtilis, Streptomyces and

Pseudomonas (Berg 2009; Cawoy et al. 2011) (Table 1).
Thus it may be said that in the light of developing ecofriendly agricultural
approaches, biopesticides are gaining commercial acceptance at a satisfactory pace
and most successful of them are microbe based products. PGPR based control of
plant diseases can hence be seen as the future of sustainable agriculture.
4 Mechanisms of Biological Control
Various direct and indirect mechanisms are involved in control of soil borne plant
pathogens by the biocontrol agent depending upon the antagonistic organism in
question. Broadly, pathogen suppression occurs either by the retarding the sapro-
phytic growth of the pathogen thereby reducing the frequency of infections through
antagonistic activities, and/or by stimulating the ‘induced systemic resistance’ in the
host-plants (van Loon et al. 1998). A variety of substances produced by plant growth
promoting rhizobacteria like siderophores, antibiotics, other small molecules, and
various kinds of extracellular enzymes like chitinases, proteases etc. are involved in
the mechanism(s) responsible for the antagonistic activities of such organisms
against phytopathogens (Glick and Bashan 1997). It may be concluded that the
direct approaches for pathogen control include antibiosis, competition, parasitism
and degradation of pathogenicity factors of the pathogen such as toxins biosurfac-
tants (Berg 2009) assisted by indirect mechanisms like improvement of plant nutri-
tion, damage compensation, changes in root system anatomy, and activation of plant
defense mechanisms, leading to enhanced plant resistance (Barea et al. 2005).
Another indirect mechanism of biological control may be plant growth promotion
by PGPR as growth promotion of plant results in shortening the time period for
which a plant is in a susceptible state, thereby leading to disease escape e.g. in the
case where PGPR cause enhanced seedling emergence rate, thereby reducing the
susceptible time for pre emergence damping-off (Kloepper et al. 1999) (Fig. 2).
4.1 Antibiosis
Antibiosis refers to the production of such metabolic agents by the organisms, which
have harmful effects on the others (Jackson 1965). Soil microorganisms produce
substances like diffusible antibiotics, volatile organic compounds (VOC), toxins,
and biosurfactants (Berg 2009) which inhibit the pathogens in the rhizosphere.
Antibiotics are extracellular secondary metabolites that are effective at low concen-
trations. Antibiotics are highly specific in their target range, have a systemic action
and are able to affect the pathogen even when they are deeply located inside the
plant tissue because they are absorbed by the plant or the seed (Dekker 1963). Apart
from that antibiotics are not easily washed away by rain and thus are effective for
long periods. Production and activity of antibiotics in the soil is affected by number
324
Table 1 Biocontrol of fungal pathogens by plant growth promoting rhizobacteria (PGPR). Note the variety of PGPR strains involved as well as the mecha-
nisms exhibited by them for inhibiting the pathogen. Also observe the range of plants for which disease control has been achieved
Percentage disease
Biocontrol agent Target pathogen Plant/disease Mechanism of action control Reference
B. subtilis Phytophthora Phytophthora blight Siderophores, HCN, 86.8% Lee et al. 2008
capsici (Red pepper) IAA, phosphatase
and ACC-deaminase
Rhizobium meliloti Macrophomina Charcoal rot Siderophores, IAA Arora et al. 2001
phaseolina (Groundnut)
Trichoderma viride Pyrenochaeta Corky root (Tomato) Competition, 81.2% Fiume and Fiume 2008
Streptomyces spp. AtB42 lycopersici mycoparasitism 75.6%
Bacillus subtilis M51 PI antibiotics, antifungal 66.8%
metabolites and
Bacillus subtilis Fusarium Wilt (Tomato) enzymes 61.1% Adebayo and Ekpo
oxysporum 2004
Pseudomonas chlorora- Fusarium Foot and root rot Phenazine-1-carboxamide, 57.7% Chin-A-Woeng et al.
phis PCL1391 oxysporum (Tomato) HCN, chitinases and 1998
proteases.
Talaromyces flavus Verticillium Verticillium wilt Naraghi et al. 2010
albo-atrum (Tomato)
Streptomyces griseus F. oxysporum f. sp. Wilt (Tomato) Chitinases 79.3% Anitha and Rabeeth
lycopersici 2009
HCN hydrogen cyanide, IAA inodole acetic acid, ACC 1-aminocyclopropane-1-carboxylic acid
P. Prashar et al.
Biological control by PGPR
Directly by inhibiting Indirectly by inducing

the growth of Indirectly by improvement
systemic resistance in
pathogen of plant nutrition and growth
the host - plant
Secretion of Competition Parasitism of

antagonistic among pathogen by Salicylic Salicylic
substances biocontrol biocontrol acid acid
like agent and agent dependent independent
antibiotics pathogen for
and various nutrients and
kinds of niches in
enzymes rhizosphere
Fig. 2 Mechanisms of biocontrol by PGPR. Inhibition of the pathogen may be achieved by any
one of these mechanisms or generally by synergistic effect of two or more mechanisms. PGPR-
plant growth promoting rhizobacteria
of physical factors like temperature, (Shanahan et al. 1992), moisture content

(Georgakopoulos et al. 1994), pH (Chin-A-Woeng et al. 1998; de Souza et al. 2003)
and nutrient sources (Duffy and Défago 1999) (Fig. 3).
Worldwide, a number of research groups are involved to isolate and exploit anti-
biotic compounds produced by rhizobacteria for control of phytopathogens. Huge
volume of literature, supporting the role of antibiotics in suppressing the plant patho-
gens and their corresponding diseases has been generated in last five decades. Many
strains of fluorescent Pseudomonas have been reported as suppressive to phytopatho-
genic fungi due to their antibiotic producing capacity along with other traits. Well-
characterized antibiotics of Pseudomonas with biocontrol properties include
phenazines, 2, 4-diacetylphloroglucinol, pyoluteorin, pyrrolnitrin, lipopeptides, and
hydrogen cyanide (Nowak-Thompson et al. 1994; Haas and Keel 2003). Another
important group of PGPR involved in antibiotic production is Bacillus species which
produces more than two dozen antibiotics having a variety of structures (Stein 2005).
Apart from Pseudomonas and Bacillus, several other bacterial strains like Serratia,
Agrobacterium and Streptomyces are also known to produce antibiotics affecting
plant pathogens. Studies using various mutant strains incapable of producing antibi-
otics like phenazines (Thomashow and Weller 1988; Hamdan et al. 1991), 2,
4-diacetylphloroglucinol (Shanahan et al. 1992) or phloroglucinols (Keel et al. 1992)
have shown that the disease suppressing capabilities of such mutants were significantly
less compared to the corresponding wild-type and complemented mutant strains
while having the similar root-colonizing capacities. At the same time cloning with
the genes responsible for antibiotic synthesis like 2, 4-diacetylphloroglucinol in non-
producing strains (Fenton et al. 1992) or introduction of multiple copies of genes for
Antibiosis
PGPR
PGPR
PGPR PGPR
PGP
R
PG Pathogen
PR c
bioti
Anti Antibioti
c
Antibiotic Pathogen
Pathogen
io tic
Antib
Pathogen Pathogen
Pathogen
Fig. 3 Antibiosis. It refers to the secretion of certain molecules e.g. antibiotics, by an organism,
which have inhibitory effects on the others. Plant growth promoting rhizobacteria produce huge
variety of diffusible and volatile antibiotics in the rhizosphere which prevent the growth of patho-
gens even when they are located deep inside the plant tissue and are effective at low concentrations
as well. PGPR: plant growth promoting rhizobacteria
antibiotics like phenazine (Fernando and Pierson 1999) resulted in enhanced antagonism
towards various pathogens. Genetic systems involved in the biosynthesis and regula-
tion of many of these antibiotics have also been well understood (Laville et al. 1992;
Raaijmakers et al. 1997; Nowak-Thompson et al. 1994; Delany et al. 2000; Mavrodi
et al. 2001). A summary of different kinds of antibiotics produced by various PGPR
alongside their target pathogen is given in Tables 2, 3, 4, and 5.
Though antibiotics have proved to be highly successful in controlling a range of
plant pathogens, in-vitro, their efficiency under actual field conditions is variable. It
has been found that quantity of antibiotics produced under field conditions in the
rhizosphere are below the minimal inhibitory concentration required for the sup-
pression of plant pathogens (Fernando et al. 2005). This may be due to various fac-
tors like the biotic and abiotic complexity of the soil, the irreversible binding of the
antibiotic to soil colloids or organic matter or microbial degradation. Factors like
chemotaxis, motility, interspecies signaling, adhesion, secretion, ability to compete
for nutrients with indigenous microbial populations within the rhizosphere and
stress response of the bacteria effect the antifungal action of the antibiotic producers
(Walsh et al. 2001). Hence it may be concluded that antibiotics at very low concen-
tration may induce systemic resistance either directly or due to their interaction with
other extra cellular metabolites that may trigger induced systemic resistance
(Fernando et al. 2005). So it is very essential to understand various factors affecting
Table 2 Antibiotics produced by the genus Pseudomonas. Note the huge variety of antibiotics produced as well as the range of plants and pathogens for which
these are effective. Also note that same antibiotic may be produced by different starins and it may affect more than one kind of pathogen, as well
Antibiotic Source organism Target Pathogen Disease Reference
2,4-diacetylphlorogluci- P. fluorescens CHA0 Thielaviopsis basicola Black root rot (Tobacco) Keel et al. 1990
nol (DAPG) P. fluorescens CHA0 Gaeumannomyces graminis Take-all disease (Wheat) Keel et al. 1992
var. tritici
P. fluorescens 05 Pyricularia oryzae and Blast and Sheath blight (Rice) Battu and Reddy 2009
Rhizoctonia solani
P. fluorescens F113 Erwinia carotovora Soft rot (Potato) Cronin et al. 1997
P. aureofaciens Q2-87 Gaeumannomyces graminis Take-all disease (Wheat) Vincent et al. 1991
var. tritici
Pyrrolnitrin P. cepacia 5.5B Rhizoctonia solani Stem rot (Poinsettia) Cartwright et al. 1995
Phenazine P. fluorescens 2–79 Gaeumannomyces graminis Take-all disease (Wheat) Gurusiddaiah et al. 1986
var. tritici and Pythium
P. chlororaphis GP72 Alternaria solani Liu et al. 2007
P. aureofaciens Rhizoctonia solani Pierson and Pierson 1996
Pyoluteorin P. fluorescens CHA0 Pythium ultimum Damping-off (cress) Maurhofer et al. 1994
P. fluorescens Pf-5 Pythium ultimum Damping-off Howell and Stipanovic 1980
Aerugine P. fluorescens Pf1TZ Botrytis cinerea Cotton and Vine Kilani-Feki et al. 2010
P. fluorescens MM-B16 Colletotrichum orbiculare Phytophthora (Pepper) Lee et al. 2003
Phytophthora capsici Anthracnose (Cucumber)
Phenazine-1- P. chlororaphis Fusarium oxysporum f. sp. Foot and root rot (Tomato) Chin-A-Woeng et al. 1998
Biocontrol of Plant Pathogens Using Plant Growth Promoting Bacteria
carboxamide radicis-lycopersici
Phenazine-1-carboxylic P. fluorescens and Gaeumannomyces graminis Take-all disease (Wheat) Thomashow et al. 1990
acid P. aureofaciens 30-84 var. tritici
Cepacidine A P. cepacia AF2001 P. ultimum Cotton and Cucumber Lee et al. 2000
Viscosinamide P. fluorescens DR54 Pythium ultimum and Nielsen et al. 1999
Rhizoctonia solani
(continued)
327
Table 2 (continued)
328
Antibiotic Source organism Target Pathogen Disease Reference

Cyclic lipopeptides P. fluorescens 96.578 Rhizoctonia solani Nielsen et al. 2000
(tensin, amphisin) P. fluorescens DSS73 Rhizoctonia solani Koch et al. 2002
N-butylbenzene Pseudomonas sp. AB2 P. ultimum, P. capsici, Inouye et al. 2000
sulphonamide R. solani, and B. cinerea
Pyocyanine P. aeruginosa Sha8 Candida albicans Hassanein et al. 2009
Hydrogen cyanide P. fluorescens CHA0 Thielaviopsis basicola Black root rot (Tobacco) Weller and Thomashow
1993
P. Prashar et al.
Table 3 Antibiotics produced by the genus Bacillus. Note the variety of antibiotics produced is lesser in comparison to that of Pseudomonas. Same antibiotic
may affect more than one pathogen
Antibiotic Source organism Target pathogen Disease Reference
Zwittermycin A B. cereus UW85 Pythium aphanidermatum Fruit rot (Cucumber) Smith et al. 1993
Phytophthora medicaginis Damping off (Alfa Alfa) Silo-Suh et al. 1994
Cyclic lipopeptides (Fengycin, B. subtilis Fusarium graminearum Anthracnose (Pepper) Wang et al. 2007
Surfactin) B. thuringiensis CMB26 Colletotrichum gloeosporioides Kim et al. 2004
B. subtilis RB14 Rhizoctonia solani Damping-off (tomato) Asaka and Shoda 1996
B. amyloliquefaciens Colletotrichum dematium, Anthracnose (Mulberry) Yoshida et al. 2001
B. subtilis Gibberella zeae Head blight (Wheat) Dunlop et al. 2011
Kanosamine B. cereus UW85 Phytophthora medicaginis Damping-off (Alfa Alfa) Milner et al. 1996
Peptide antibiotics (Gramicidin Bacillus brevis Botrytis cinerea Grey mold (Strawberry) Haggag 2008
S and Polymyxin B)
329
330
Table 4 Antibiotics produced by genus Streptomyces. The variety of antibiotics produced is much lesser in comparison to that of genus Pseudomonas. Note
that same antibiotic may affect more than one kind of pathogen
Phoslactomycine I Streptomyces nigrescens Botrytis cinerea and Alternaria Fushimi et al. 1989
kikuchiana
Phenylacetic acid and sodium Streptomyces humidus Pythium ultimum, Phytophthora Phytophthora Hwang et al. 2001
phenylacetate capsici, Rhizoctonia solani blight (Pepper)
Tubercidin Streptomyces violaceoniger A50 Phytophthora capsici Magnaporthe grisea Hwang et al. 1994
Macrolide antibiotic (Oomycin A) Streptomyces libani Phytophthora capsici Anthracnose, and Kim et al. 1999
leaf blast
Aerugine Streptomyces fradiae SU-1 Colletotrichum gloeosporioides and Apichaisataienchote
Phytophthora parasitica et al. 2006
P. Prashar et al.
Table 5 Antibiotics produced by genus Serratia. Note the target organism is mainly Sclerotinia sclerotiorum comapred to broad range of target organism in
case of other rhizobacteria. Also fewer kinds of antibiotics are produced by Serratia as compared to other rhizobacteria
Pyrrolnitrin S. plymuthica IC14 Botrytis cinerea and Sclerotinia Gray and white mold disease Kamensky et al. 2003
sclerotiorum (Cucumber)
Macrolides: Haterumalide A S. plymuthica Sclerotinia sclerotiorum Suppression of apothecial formation Thaning et al. 2001
and ascospore germination.
Haterumalide NA, B, NE, S. plymuthica A 153 Sclerotinia sclerotiorum Suppression of apothecial formation in Levenfors et al. 2004
and X sclerotia
331
Competition among biocontrol agent and target pathogen
Plant derived nutrients
Niches Resources Iron
Flagellar motility Growth factors
Traits
Chemotaxis Catabolism of root exudates
Traits
Surface antigens Siderophore
Surface appendages
Agglutination
Phenotypic diversity
Antibiotic Production
Fig. 4 Competition among biocontrol agent and target pathogen. It occurs for suitable niches and
available resources in the rhizosphere. Possession of traits listed in the figure help the biocontrol
agent to out-compete the pathogen thereby controlling the corresponding disease
the production and activity of antibiotics along with their interaction with other
metabolites to induce systemic resistance in plants, before we can fully utilize the
potential of antibiotics in disease management.
4.2 Competition
Competition is the active demand in excess of immediate. Given the fact that soils
generally have limited and inaccessible forms of nutrients for microbial utilization;
competition for resources such as nutrients, oxygen and suitable sites at the root
surfaces is a commonly occurring phenomenon among soil inhabiting organisms
(Cawoy et al. 2011) and is a fundamental mechanism for the control of soil borne
pathogens. Starvation is an important and common cause of microbial death, so
competition for limiting nutrients, may lead to biological control of fungal phyto-
pathogens (Eisendle et al. 2004). However competition will take place only if the
introduced microbial strain and the pathogen have similar demands in terms of
space and resources (van Dijk and Nelson 2000). Since microbes generally have
common requirements in terms of essential growth factors, lack of critical nutrients,
like carbon (Alabouvette et al. 2006) and mineral elements, like iron, along with
suitable sites in the rhizosphere, is assumed to be responsible for inhibition of fun-
gal spore germination in soil (Loper and Henkels 1997) (Fig. 4).
4.2.1 Competition for Root Niches
Availability of carbon is the limiting factor for microbial activity in soil and thus
affects a number of processes mediated by soil microflora (Paterson 2003). Since the
root surface itself and its immediate surroundings have high carbon concentrations
(equivalent to 15–60% of the total photosynthetic production of the plant) owing to
rhizodeposition (Curl and Truelove 1986; Lynch and Whipps 1990), there is an acute
competition among rhizospheric microorganism to occupy suitable nutrient rich sites
along the root surface (Compant et al. 2005). Traits like flagellar motility of PGPR
strains such as (de Weger et al. 1987a), chemotaxis toward root exudates and surface
antigen (O-antigen of lipopolysaccharide) mediated attachment to the root surface,
play crucial role in successfully establishing the introduced strain in the rhizosphere
(de Weger et al. 1987b; Tilak et al. 1999; de Weert and Bloomberg 2006). Various
studies involving the use of mutant strains lacking in above mentioned capabilities
have further confirmed their significant role in the root colonization. For example
nonmotile mutants of P. fluorescens WCS365 have been proved to have reduced root
colonization in tomato, wheat, and radish crop (Dekkers et al. 1998a). Similarly
Capdevila et al. (2004) examined the non-motile mutants of a well-known biocontrol
strain Pseudomonas fluorescens F113 for competitive root colonization. It was
observed that all the mutants were completely displaced from the root tip when com-
peting with the wild-type F113 strain. Moreover one of the mutants which had longer
flagella compared to the parent strain, when grown in rich medium showed a higher
motility, suggesting the possibility of improving competitive root colonization by
manipulating the motility processes. de Weert et al. (2002) established the indepen-
dent role of chemotaxis along with motility. They evaluated cheA mutants of
Pseudomonas fluorescens that were defective in flagella-driven chemotaxis but still
having motility for competitive tomato root-tip colonization in a gnotobiotic sand
system and in nonsterile potting soil. Mutants were found to have a strongly reduced
competitive root colonizing ability after seven days of plant growth, in both the cases.
Substances that are known to create chemotactic response toward root exudates
include organic acids, amino acids, sugars etc. (de Weert et al. 2002).
Same way, the mutant studies have proved that the presence of O-antigen
lipopolysaccharide (de Weger et al. 1989a) and ability to synthesize amino acids
(Simons et al. 1997) and vitamin B1 (Simons et al. 1996) are important root coloni-
zation traits. However the role of O-antigen lipopolysaccharide may vary with
strain. As de Weger et al. (1989b) reported that in mutants of strains WCS358 and
WCS374 which lacked the 0-antigenic side chain of the lipopolysaccharide, the
adhesion to potato roots was not effected, but the mutant of P. fluorescens PCL1205
which had shorter O-antigen chain was defective in colonization ability (Dekkers
et al. 1998b).
Other cell surface exopolysaccharides, like cellulose fibrils, also play a role in
the initial adherence of rhizobacteria like A. tumefaciens and R. legunisprum to
plant surface and root hairs (Smit et al. 1986; Weller 1988). Extracellular fibrils
produced by R. leguminosarum were found to have a positive correlation between
the percentage of fibrilated cells and the ability of the bacteria to form caps and to
adhere to pea root hair tips. Laus et al. (2005) further confirmed this role by their
mutagen studies. They reported that wild-type Rhizobium leguminosarum RBL5523
infected the elongated growing root hairs whereas its cellulose fibril-deficient celE
mutant infected young emerging root hairs. Further, exopolysaccharide-deficient
strains that retained the ability to produce cellulose fibrils could also infect elon-
gated root hairs but infection thread colonization was defective. Rhizospheric bac-
teria like Rhizobium, Azospirillum, Pseudomonas and nitrogen-fixing endophytes
are known to embed and encase themselves in geltinaceous extracellular polysac-
charide material, just like in other ecological niches, leading to the formation of
biofilms and hence may escape from displacement by indigenous microflora in the
root zone (Weller 1988; Rodriguez-Navarro et al. 2007; Meneses et al. 2011).
Sporulation has also been found to influence root colonization in Bacillus spp.
(Zheng and Sinclair 2000).
Bacterial surface appendages like fimbriae also mediate the attachment of the
introduced strain to the root surfaces (Vesper and Bauer 1986; Weller 1988). These
proteinaceous, fibrillar structures are generally found on the surfaces of many gram-
negative bacteria and play a role in attachment of the bacterium to different surfaces
(Weller 1988). Vesper and Bauer (1986) analyzed the role of pili (fimbriae) in
attachment of bacteria to soybean roots, by using five strains of Bradyrhizobium
japonicum and one strain of Rhizobium trifolii. It was observed that there was cor-
relation between the total number cells that colonized the roots and the cells pos-
sessing pili. Role of fimbriae has also been established in case of Pseudomonas.
Vesper (1987) observed a similar correlation in P. fluorescens 2–79 for attachment
to corn roots. It was found that the nonmucoidal strain 2–79 was highly piliated
whereas the mucoidal type was nearly devoid of pili and the presence of pili was
quantitatively correlated with hydrophobic attachment to roots.
One more factor that has been shown to be instrumental in early steps of root colo-
nization i.e. attachment of the bacterial cells to the root surface, is agglutination.
Bacterial cells may be agglutinated by root surface mucilaginous substances like
pectic compounds, other acidic polysaccharides (Anderson and Jasalavich 1979;
Slusarenko et al. 1983) and certain unknown compounds present in the root exudates
(Anderson 1983; Chao et al. 1988). It has been shown that the agglutination and
adherence to root surface leading to the long term colonization is rapid and frequent
in rhizosphere bacteria like P. fluorescens than that of the non-rhizospheric bacteria
in various plants like radish (James et al. 1985) and pea (Chao et al. 1988). Certain
strains of Pseudomonas like P. putida and P. fluorescens have been shown to get
agglutinated by a glycoprotein complex, termed agglutinin, released from root sur-
faces of plant (Anderson et al. 1988). Mutants of P. putida, lacking the ability to
agglutinate with components present in the washes of bean and cucumber were com-
pared with their Agg + parental strain for their capacity to colonize the roots of cucum-
ber plants and control Fusarium oxysporum f. sp. cucumerinum causing wilt. All the
mutants showed limited root colonization as well as disease control compared to the
parent strain (Tari and Anderson 1988). Thus agglutination properties of PGPR
may give them an edge over the pathogens in successfully occupying the suitable
sites and hence outgrow in the rhizosphere. However all PGPR may not have similar
agglutination potential. For example various strains of fluorescent Pseudomonas

have been shown to behave differently to the presence of agglutinins (Jasalavich and
Anderson 1981) and the root agglutinins may not always play a decisive role in root
colonization though they may be involved in short-term adherence of Pseudomonas
to roots (Glandorf et al. 1994). In their study Glandorf et al. did not observe any cor-
relation between the potato root colonization by rhizospheric Pseudomonas and their
agglutination ability, with both the low and high molecular-weight agglutinins.
Moreover, no difference in potato root colonization was observed for Agg- mutants
of P. putida colonization and their parental strain. In another study by Glandorf et al.
(1993) crop-specific agglutination of rhizospheric fluorescent Pseudomonas spp. for
potato, grass and wheat was investigated. It was found that the composition of rhizo-
sphere Pseudomonas populations varied with crops, but crop-specific root coloniza-
tion or crop-specific agglutination was not observed.
Another determinant of root colonization potential is the ability of introduced
strain to generate phenotypically diverse population by site-specific recombinases
of the one integrase family which control the phase variation (Weller 2007). Gene
sss and xerD have been found to encode two different types of site-specific recom-
binases that determine phenotypic variation during rhizosphere colonization
(Dekkers et al. 1998c; Sanchez-Contreras et al. 2002; Martınez-Granero et al. 2005).
A mutant of an efficient root colonizer P. fluorescens WCS365 with impaired colo-
nization abilities in potato, radish, wheat, and tomato was found by Dekkers et al.
(1998c). This mutant strain (PCL1233) was found to lack site-specific recombi-
nases encoded by the sss gene, which play a role in phase variation. Similarly
mutants of P. fluorescens F113 affected in the sss or xerD genes showed significantly
low quantity of phenotypic variants compared to the wild-type strain and were
found to be severely impaired in competitive root colonization(Martınez-Granero
et al. 2005). Root exudates are assumed to affect the expression of genes site-specific
recombinases. Further it has been shown that introduction of the extra copies of site-
specific recombinase gene and its transfer to a rhizosphere-incompetent Pseudomonas
strain improves the rhizosphere colonization (Dekkers et al. 2000) as well as their
biocontrol abilities (Chin-a-Woeng et al. 2000).
Production of antibiotics also influences the ecological competence of the pro-
ducing strain in terms of root colonization and hence their persistence in the rhizo-
sphere leading to beneficial interaction with plants by stimulating host defense
mechanisms (Ongena and Jacques 2008). For example phenazines produced by two
strains of Pseudomonas which are known to suppress take-all of wheat caused by
Gaeumannomyces graminis var. tritici, i.e. P. fluorescens 2–79 and P. aureofaciens
30–84, have been also shown to contribute to the ecological competence of these
strains in the rhizosphere of wheat (Mazzola et al. 1992). Mutants defective in
phenazine production were found to have significantly reduced population size
compared to the parental strains. Similarly lipopeptides antibiotics: surfactins,
iturins and fengycins produced by Bacillus species enhance their competitive poten-
tial. The ability to utilize nutrients like vitamins biotin; thiamine (Streit et al. 1996)
and amino acids like aspartic acid; glutamic acid; iso-leucine; leucine and lysine
(Simons et al. 1997) also plays a crucial role in root colonization by the PGPR.
Abiotic factors like oxygen supply, temperature, pH, water content and soil type
also tend to affect the process of colonization (Weller 1988). Thus, in order to out-
compete the indigenous microflora, including the pathogens, successful root coloni-
zation is a prerequisite for the introduced biocontrol agent.
4.2.2 Competition for Iron
Iron is an essential nutrient for all living organisms (Neilands 1981). Though pres-
ent abundantly on earth (fourth largest mineral), it is not readily available to microbes
and plants as it get easily oxidized and exist predominantly in the form of sparingly
soluble ferric ions which cannot be utilized directly (Neilands et al. 1987). The
concentration of iron in the soil may be as low as 10−18 M which is significantly
lesser than the required concentrations of approximately 10−6 M for the growth of
microorganisms (Pal and Gardener 2006). Soil microorganisms are known to secrete
a class of low molecular weight molecules (400–1,000 Da) , known as siderophores,
which act as ligands for binding ferric ions due to their high iron affinity (Kd = 10−20
to 10−50) (Castignetti and Smarrelli 1986). Thus siderophore producing strains have
a selective advantage over the non producing strains, including pathogens, as they
may competitively scavenge the limited amount of ferric ions available in the rhizo-
sphere resulting in inhibition of growth of the pathogens in their immediate vicinity
due to iron limitation (O’Sullivan and O’Gara 1992; Haas and Défago 2005).
A wide range of PGPR are known to produce siderphores including Pseudomonas
(Kloepper et al. 1980; Elad and Baker 1985), Bacillus (Grossman et al. 1993;
Chaiharn et al. 2009; Yu et al. 2011), Acinetobacter (Sarode et al. 2009), Serratia
(Kamensky et al. 2003; Berg et al. 2005) etc. and hence this serves as an important
phenomenon contributing to suppression of soil borne pathogens (Fig. 5).
Certain fungal phytopathogens may also synthesize siderophores, however their
affinity for iron is much lower compared to those of antagonistic PGPR (Schippers
et al. 1987) and as a result PGPR out-compete fungal phytopathogens for the avail-
able iron. Overall, soil borne pathogens whose route of infection is through myce-
lial contact, such as species of Fusarium and Pythium, are more susceptible to face
competition from other soil and plant-associated microbes than those pathogens
that germinate directly on plant surfaces and infect through appressoria and infec-
tion pegs (Pal and Gardener 2006). Siderophores are generally classified on the
basis of the ligand that they use for chelation of ferric ions, as either catecholates
(phenolates) or hydroxamates or carboxylates (Ankenbauer et al. 1988; Winkelmann
and Drechsel 1997). While phenolate siderophores are produced exclusively by
bacteria, hydroxamates are produced by both bacteria and fungi (Cox et al. 1981).
Pseudomonas generally produce two kinds of siderophores: pyoverdins which are
yellow-green siderophores produced by fluorescent Pseudomonas in iron-limited
conditions (Hamdan et al. 1991) and pyochelin which is a salicylate derivative and
structurally unique phenolate siderophore produced by P. aeruginosa (Cox et al.
1981). Role of siderophores in control of pathogens either singly or synergistically
with other metabolites like antibiotics has been well established, particularly by
analyzing the mutants deficient in siderophore production (Loper 1988; Buysens
Competition for Iron
Pseudomonas
Pseudomonas Pathogen
Pathogen
Siderphore
Siderphore
Fe3+
Fe3+
3+
Fe
Fe3+ Fe3+
Fig. 5 Competition for iron. Iron is an essential growth requirement of microbes. Most of the
plant growth promoting rhizobacteria like Pseudomonas, release small ligands called siderphores
for binding ferric ions available in the rhizosphere, as they cannot be utilized directly. This results
in little or no ferric ions left for pathogen use and thus their inhibition due to iron scarcity. Fe 3+:
Ferric ion
et al. 1994). Numerous studies have reported the role of siderophore in suppression
of plant diseases caused by Pythium spp., Fusarium oxysporum, Magnaporthe gri-
sea, Rhizoctonia solani, Pyricularia oryzae, Alternaria, Sclerotium sp. and many
other fungal pathogens in plants like wheat, tomato, rice, chick-pea, green gram by
Pseudomonas, Streptomyces sp., Alcaligenes strain MFA1 (Kloepper et al. 1980;
Elad and Baker 1985; Yuen and Schroth 1986; Kumar and Dube 1992; Buysens
et al. 1994; Chaiharn et al. 2009; Reddy et al. 2010; Sahu and Sindhu 2011). Disease
suppressing capability of siderophores is affected by factors like the plant, target phyto-
pathogen, soil properties, siderophore producing strain and type of siderophore
(Glick and Bashan 1997). However siderophores do not always contribute to the
biocontrol potential of the rhizobacteria. For example P. fluorescens 2–79 which
is a well known strain for suppressing take-all disease of wheat caused by
Gaeumannomyces graminis var. tritici is known to produce phenazine-1-carboxylic
acid and a fluorescent pyoverdin siderophore (Thomashow and Weller 1988).
A study was designed to investigate the role of pyoverdin siderophores in disease
suppression. Mutants of P. fluorescens 2–79, deficient in production of the antibiotic,
the siderophore, or both factors were compared with the parental strain for control
of take-all on wheat roots. It was observed that pyoverdin-deficient mutants were
equally effective in controlling the disease as their respective fluorescent parental
strains (Hamdan et al. 1991). Further it was observed that under iron limitation cer-
tain other substances distinct from pyoverdin siderophores were produced which
facilitated iron uptake and fungal inhibition. Thus these observations still support the
role of iron competition though mediated by molecules other then siderophores.
Another interesting fact is that pathogen suppression under iron limited conditions
by pyoverdin siderophore producing bacteria is more effective at alkaline and neutral
soils as compared to acidic soils. The possible explanation for this kind of behavior
is increasing solubility of Fe3+ species with decreasing pH (Weller 1988; Haas and
Défago 2005). The second kind of siderophore produced by , pyochelin has recently
been reported to exhibit moderate affinity for Fe3+ (pFe = 16.0 at pH 7.4) (Brandel
et al. 2012) in contrast to earlier reports that marked pyochelin as having very low
iron-binding coefficient, 5 × 105 (Cox et al. 1981; Visca et al. 1992). Moreover it is a
strong chelator of divalent metals such as Zn (II) (pZn = 11.8 at pH 7.4) and Cu (II)
(pCu = 14.9 at pH 7.4) (Brandel et al. 2012) and bind other metals like Co (II), Mo
(VI) and Ni (II), with appreciable affinity (Visca et al. 1992). The role of pyochelin
siderophore in the biocontrol Pythium splendens induced damping-off in tomato,
along with pyoverdin has been confirmed in P. aeruginosa 7NSK2 by using pyoche-
lin – deficient mutants derived from P. aeruginosa 7NSK2 (Buysens et al. 1996).
Further, the effect of two siderophores i.e. pyoverdin and pyochelin was assumed to
be mutually exchangeable because a mutant producing only pyochelin and not
pyoverdin was equally antagonistic as a mutant producing only pyoverdin and not
pyochelin. In another study, a novel species of thermophilic Pseudomonas was found
to produce pyochelin with antifungal properties against several Candida species and
Aspergillus fumigatus (Phoebe et al. 2001). Hence, it may be concluded that compe-
tition for iron and other mineral ions like Cu2+ and Zn2+ is one of the significant
mechanism followed by PGPR to suppress the soil borne pathogens, however sidero-
phores alone may not be sufficient to suppress the disease in all cases.
4.2.3 Competition for Plant Derived Nutrients
Many studies have reported competition for root exudates and other nutrients among
rhizosphere bacteria as the mechanism utilized by PGPR for biocontrol of patho-
gens. For example Elad and Chet (1987) evaluated P. putida or P. cepacia for the
control of Pythium aphanidermatum in wheat, tomato, cucumber, melon, bean, pep-
per, tomato and cotton plants. It was found that competition for nutrients between
germinating oospores of P. aphanidermatum and bacteria, was responsible for sup-
pression of oospore germination and hence disease control up to 60–90%. The pop-
ulation of P. putida or P. cepacia was much higher compared to other bacteria and
it retarded the establishment of Pythium along the roots, which are the potential
sites for its attack. Further, the production of inhibitory substances, antibiotics and
lytic enzymes was not observed and role of siderophores was excluded as the pH of
the soil was 6.0 which ensured the availability of iron in high concentration. In a
similar study competition for plant-derived unsaturated long-chain fatty acids was
reported to be the mechanism of biocontrol of the seed-rotting oomycete, Pythium
ultimum in cotton seedling, by Enterobacter cloacae, as fatty acids from seeds and
roots are essential for germination responses of P. ultimum (van Dijk and Nelson
2000). As in case of siderophores, the use of mutants has confirmed that competi-
tion for nutrients is a major contributing factor in the control of soil borne pathogens
(van Dijk and Nelson 2000). Another essential requirement for the successful estab-
lishment of the introduced microbial strain in the rhizosphere is the effective catabo-
lism and utilization of the root exudates so that they are not available for the pathogen
utilization. For example biocontrol of Pythium damping off by E. cloacae in various
tested seeds like carrot, cotton, cucumber, lettuce, radish, tomato, wheat, corn and
pea was found to be dependent on its ability to metabolize long-chain fatty acids,
mainly linoleic acid, released from the seed during germination so as to reduce their
availability to P. ultimum sporangia (Kageyama and Nelson 2003). It was observed
that P. ultimum was suppressed in all cases except in corn and pea seeds. The reason
for this is based on the types of root exudates released by these two plants. Both,
corn and pea seeds release high levels of simple sugars during early stages of seed
germination which repress b-oxidation, thereby preventing the metabolism of exu-
dates fatty acids. In such conditions E. cloacae predominantly metabolizes simple
sugars than fatty acids, which then induce the sporangium germination. Hence, it
may be said that while competitive uptake of root exudates by the biocontrol agent
results in scavenging of available nutrients and growth factors for the pathogen, it
may vary according to the type of exudates (Fig. 6).
Competition for plant derived nutrients
Seed and
Germinating seed
root Seed and

exudates root
exudates
Seed and
root
exudates
PGPR
Pathogen PGPR
Pathogen PGPR
PGPR
Pathogen
PGPR
No spore germination Pathogen inhibition
Fig. 6 Competition for plant derived nutrients. A number of seed and roots exudates released in
rhizosphere e.g. fatty acids like linoleic acid, are required for the germenation of spores of fungal
pathogen in soil. Plant growth promoting rhizobacteria usually hold higher affinity and effective
catabolic capacity for utilizing such exudates and hence scvange the rhizosphere of essential exu-
dates required by pathogens thereby inhibiting them
4.3 Parasitism
It is an important mechanism employed by certain PGPR, actinomycetes in particu-

lar, to suppress the pathogens, but is more commonly used by fungal strains like
Trichoderma (Haran et al. 1996; Whipps 2001; Viterbo et al. 2002). This kind of
control requires close contact between the target (fungal pathogen) and the host to
ensure target recognition (chemical gradients and mycelial surface features), attack,
penetration and subsequent lyses and degradation of fungal spores or hyphae
(Handelsman and Stabb 1996; Benitez et al. 2004). Rhizospheric bacteria recognize
fungus in various morphological forms including spores; hyphae; fruiting bodies
and their attachment is affected by flagella mediated chemotaxis towards mycotox-
ins and other fungal metabolites (Hogan et al. 2007). The degree of attachment may
range from simple attachment of the parasite to the host (Nelson et al. 1986;
Wisniewski et al. 1989) to the engulfment of the fungal pathogen in biofilm induc-
ing the lysis of the fungal cells (Hogan et al. 2007). The most common bacterial
strains that are known as fungal colonizers include Pseudomonas and Burkholderia
(Hogan et al. 2007). Extensive colonization of the fungal hyphae by P. fluorescens
WCS365 and P. chlororaphis PCL1391 was observed as a key factor in the biocontrol
of foot and root rot of tomato plants caused by Fusarium oxysporum (Bolwerk et al.
2003). Lysis of the target fungal pathogen is generally caused by destruction of the
cell wall by extracellular hydrolytic enzymes including chitinases, proteases and
glucanases (Chet 1987). Though, literature reports more investigations on fungal
lytic enzymes, several bacterial species like Pseudomonas (Nandakumar et al.
2007), Bacillus (Wang et al. 2002; Chang et al. 2003; Huang et al. 2005), Serratia
(Chet et al. 1990) and others are known to suppress fungal pathogens of plants
through lytic activities of chitinases (Wang and Chang 1997; Wang et al. 1999;
Radjacommare et al. 2010; Hariprasad et al. 2011) glucanases (Fridlender et al.
1993) and proteases (Dunne et al. 1997). For example Chet et al. (1990) isolated a
chitinases producing Serratia marcescens which proved to be an efficient biocontrol
agent of Sclerotium rolfsii and solani under greenhouse conditions. Further, they
cloned E. coli with the gene coding for chitinase and the partially purified chitinase
produced by the cloned gene was found to be effective in reducing disease incidence
caused by S. rolfsii in beans and R. solani in cotton, under greenhouse conditions.
Similarly three glucanase-producing endophytic actinomycetes, Actinoplanes cam-
panulatus, Micromonospora chalcea and Streptomyces spiralis, significantly
reduced damping-off and crown and root rot of cucumber caused by Pythium
aphanidermatum. When applied as a combined treatment the effect was equivalent
to the chemical fungicide metalaxyl (El-Tarabily et al. 2009). Various Bacillus
strains like Bacillus cereus YQ 308 (Chang et al. 2003), Bacillus cereus CRS 7
(Kishore and Pande 2007), Bacillus amyloliquefaciens V656 (Wang et al. 2002) and
Bacillus cereus QQ308 (Chang et al. 2007) have been found to produce extracellu-
lar chitinase, having antifungal activities against fungal pathogens like Botrytis
cinerea, Fusarium oxysporum, Fusarium solani and Pythium ultimum in plants
like chickpea and chinese cabbage. Bacillus cereus QQ308 produced enzymes like
chitosanase and proteases along with chitinases and exhibited growth promotion as
well in Chinese cabbage. In a similar study Fridlender et al. (1993) isolated b-1, 3
glucanase-producing P. cepacia which decreased the incidence of diseases caused
by Rhizoctonia solani, Sclerotium rolfsii and Pythium ultimwn by 85%, 48% and
71%, respectively. The isolated strain was found to have no chitinolytic activity.
Various mutational studies have established the significant role played by lytic
enzymes in biocontrol activities. For example, by using Tn5 insertion mutants of a
strain of Stenotrophomonas maltophilia (producing extracellular chitinase and pro-
tease), it has shown that the biocontrol potential of this strain against Pythium ulti-
mum induced damping-off in sugar beet was conferred by the extracellular protease
while chitinases do not contribute to its antagonistic behavior (Dunne et al. 1997).
Further, mutants (W81M3 and W81M4) which have protease overproduction capac-
ity exhibited noticeably improved biocontrol potential (Dunne et al. 2000). Similarly
cloning of chitinase gene chiA of B. circulans WL-12 in strain B. subtilis F29-3 has
shown that in comparison to the B. subtilis control strain, chiA expressing B. subtilis
F29-3 exhibited a greater inhibition of spore germination of Botrytis elliptica (Chen
et al. 2004). Hence it may be concluded that pathogen colonization and parasitism
is a key mechanism adopted by many rhizobacteria to inhibit the fungal pathogen
and such organisms have an edge over others in terms of effective control of the
target pathogen (Fig. 7).
Parasitism
Attachment of the parasite to the host Engulfment of the parasite by the host
Target recognition
Spores; hyphae;
fruiting bodies
Attack
Flagella
Penetration mediated chemotaxis
Extracellular
Lyses and degradation
hydrolytic enzymes
Fig. 7 Parasitism. It requires close contact between fungal pathogen and biocontrol agent, degree
of which may vary for simple attachment to complete engulfment into the biofilm produced by
biocontrol agent. Rhizobacteria recognizes various morphological forms of fungal pathogen
including spores, hyphae and fruiting bodies. Variety of extracellular hydrolytic enzymes like chi-
tinases, proteases and glucanases are released by the rhizobacteria to cause destruction of the cell
wall leading to lysis of the target fungal pathogen
4.4 Induced Systemic Resistance
Induced resistance may be defined as the enhancement of the plant’s defensive

capacity against a broad array of pathogens and parasites, wherein the plant’s innate
defenses are potentiated against subsequent biotic challenges by a stimulus prior to
infection by a virulent plant pathogen (Bakker et al. 2003; Meziane et al. 2005).
This state of enhanced basal persistence of the plant due to an inducing agent is
called as induced systemic resistance (ISR) or systemic acquired resistance (SAR)
and it depends upon the signaling compounds (van Loon 2007). Induced systemic
resistance and systemic acquired resistance are phenotypically similar and both
result in activation of latent resistant mechanisms in the plant capable of expression
on subsequent or challenge inoculation with the pathogen (van Loon 1997).
However, to differentiate between the two the inducing agents have been used as a
criterion. Systemic resistance induced by rhizobacteria has been termed as induced
systemic resistance; while the one triggered by pathogen is called as systemic
acquired resistance (van Loon et al. 1998). Thus induced systemic resistance may
be defined as plant-mediated, broad-spectrum resistance response that is activated
by selected strains of saprophytic rhizosphere bacteria specifically the PGPR.
Systemic acquired resistance requires induction by a pathogen mediated hypersen-
sitive reaction resulting in necrosis of the tissue, while induced systemic resistance
by PGPR do not produce such kind visible symptoms on the plant (van Loon
et al. 1998). However, both induced systemic resistance and pathogen-induced sys-
temic acquired resistance render uninfected plant parts more resistant towards a
broad spectrum of pathogens (Pieterse et al. 2002) and the enhanced defensive
capacity, conferred on the plant by induced systemic resistance, reduces the rate of
disease development upon infection with a challenging pathogen, thereby produc-
ing fewer diseased plants or plants with lesser disease (van Loon 2007). Induced
resistance is observed not only at the site of the initial treatment but also in distal,
untreated plant parts (Conrath et al. 2002). Though reports on systemic acquired
resistance against virus, by tobacco mosaic virus (McKinney 1929) as well as
necrotic fungi (Hecht and Bateman 1964), and bacteria (Klement et al. 1966) have
been obtained since long, the role of PGPR in induced resistance was established
quite late. Rhizobacterium-mediated induced systemic resistance was first time
reported in the year 1991, simultaneously by van Peer et al. (1991) in carnation and by
Wei et al. (1991) in cucumber. van Peer et al. observed that susceptibility of carnation
to wilt causing Fusarium oxysporum f.sp. dianthi was reduced upon treatment of plant
roots with P. flourecens WCS417 due to induction of systemic resistance. Similarly
Wei et al. (1991) reported the protection of cucumber against foliar disease caused
by Colletotrichum orbicularecarnation. Since then, the strain P. flourecens WCS417
has been shown to induce resistance in many other plants as well, like in radish
against Fusarium oxysporum (Leeman et al. 1995a), in tomato against Fusarium
oxysporum (Duijff et al. 1998) and in bean against Colletotrichum lindemuthi-
anum (Bigirimana and Höfte 2002). In all cases the spatial separation of the
pathogen and the biocontrol strains was ensured to exclude the possibility of any
other antagonistic mechanism. Many other stains of Pseudomonas have also been
shown to induce resistance. For example Cameron et al. (1994) reported biologi-
cal induction of systemic acquired resistance in Arabidopsis thaliana by local
inoculation of a single leaf with a virulent Pseudomonas syringae pv. tomato (Pst)
carrying the avrRpt2 avirulence gene. They observed that the plants were pro-
tected against infection by other virulent pathogens including P. syringae
pv.maculicola. Other non pathogenic strains have been reported induced systemic
resistance in a variety of host plant-pathogen combinations like Pseudomonas
putida WCS358 in Arabidopsis thaliana against Pseudomonas syringae pv.
tomato and Fusarium oxysporum f. sp. raphani (van Wees et al. 1997), putida
89B-27 in cucumber against Fusarium oxysporum f. sp. cucumerinum (Liu et al.
1995) and putida 89B-61 against angular leaf-spot caused by Pseudomonas
syringae pv. Lachrymans in cucumber (Wei et al. 1996). Siderophore producing
P. aeruginosa 7NSK2 has been reported to induce resistance against Botrytis
cinerea in bean and tomato (de Meyer and Höfte 1997; Audenaert et al. 2002).
Seed, root, soil and foliar application fluorescens strains viz., PF1 and FP7 either
alone or in combination (seed + root + soil + foliar) inducted systemic resistance
against sheath blight disease causing fungus Rhizoctonia solani and hence effec-
tively reduced incidence of disease and promoted plant growth and yield under
both glasshouse and field conditions (Nandakumar et al. 2001).
A number of non-pathogenic rhizospheric Bacillus strains have also been reported
to impart induced systemic resistance against fungal diseases in a variety of plants.
A non-pathogenic, phyllospheric isolate of Bacillus mycoides (Bac J) from sugar
beet reduced the Cercospora leaf spot of sugar beet caused by Cercospora beticola
Sacc. by 38–91% by inducing systemic resistance (Bargabus-Larson et al. 2002). In
a similar study induction of systemic acquired resistance against the same fungal
pathogen i.e. Cercospora leaf spot causing Cercospora beticola Sacc. in sugar beet,
was investigated for two isolates of B. pumilus from sugar beet by the same group.
It was found that two B. pumilus isolates 203–6 and 203–7, reduced the disease
occurrence by approximately 70% (Bargabus-Larson et al. 2004). Other strains of
Bacillus that have been found to induce systemic resistance in plants include
B. pumilus INR7, B. pumilus SE34, B. subtilis GBO3, against downy mildew caus-
ing fungus Sclerospora graminicola in pearl millet (Niranjan Raj et al. 2003), B.
pumilus INR7 against angular leaf-spot caused by P. syringae pv. lachrymans in
cucumber (Wei et al. 1996), B.amyloliquefaciens 937a, B. subtilis 937b, and B.
pumilus SE34 against tomato mottle virus (ToMoV) in tomato plants (Murphy and
Zehnder 2000). Krause et al. (2003) isolated eleven strains of Bacillus spp. from
compost which were found to significantly suppress the bacterial leaf spot of radish,
caused by Xanthomonas campestris pv. armoraciae, by induction of systemic resis-
tance in the host plant. However the degree of disease suppression was significantly
less compared to that caused by Trichoderma hamatum 382. Even the effectiveness
of a mixed inoculum of T382 and the most effective rhizobacterial Bacillus strain
was less effective than T382 alone.
Combinations of two or more PGPR strains have also been evaluated in various
studies, for disease control by inducing resistance. For example, a two-strain mixture
of B. subtilis GB03 and B. amyloliquefaciens IN937 protected tomato plants against

Cucumber mosaic virus and P. syringae pv. tomato DC3000 in Arabidopsis thaliana
(Ryu et al. 2007). The involvement of salicylic acid signaling in induced resistance
was also established in this study by using NahG transgenic plants, which degrade
endogenous salicylic acid. The biopreparation mixture did not induced resistance in
these transgenic plants. Similarly Muphy et al. (2003) evaluated a combination of
this strain i.e. Bacillus subtilis GB03 with either of B. pumilus SE34, B. amyloliq-
uefaciens IN937a, B. subtilis IN937b, B. pumilus INR7 or B. pumilus T4 formulated
with chitosan to protect tomato plants against cucumber mosaic virus. All the bio-
prepration treated plants showed significant disease protection as well as growth
enhancement compared to control plants. In another study by Jetiyanon and
Kloepper (2002) mixtures of compatible PGPR strains were tested to induce sys-
temic resistance against diseases of several different plant hosts compared to indi-
vidual PGPR strains. The mixture strains used were B. amyloliquefaciens IN937a + B.
pumilus IN937b, B. pumilus IN937b + B. pumilus SE34, B. pumilus IN937b + B.
pumilus SE49, and B. pumilus T4+ B. pumilus INR7. The range of host plant and
disease tested were: bacterial wilt of tomato caused by Ralstonia solanacearum,
anthracnose of long cayenne pepper (Capsicum annuum var. acuminatum) caused
by Colletotrichum gloeosporioides, damping off in Brassica chinensis var. para-
chinensis caused by Rhizoctonia solani, and cucumber mosaic virus on cucumber
(Cucumis sativus). It was observed that the most of the compatible PGPR mixtures
induced a higher level of protection than individual strains. PGPR strains other than
Pseudomonas and Bacillus are also known to suppress various phytopathogens by
induced systemic resistance mediated mechanism.
PGPR strains like Serratia marcescens 90–166 (Liu et al. 1995), Serratia marc-
escens B2 (Someya et al. 2002) and Serratia liquefaciens MG1 (Schuhegger et al.
2006) are also known to induce resistance against wide range of pathogens like rice
blast causing Pyricularia oryzae, leaf pathogen Alternaria alternata of tomato
plants, Fusarium oxysporum f. sp.cucumerinum and angular leaf-spot causing syrin-
gae pv. Lachrymans in cucumber (Wei et al. 1996), respectively. Use of mutants
(Schuhegger et al. 2006) has further confirmed the role of these PGPR in generating
resistance against target pathogens as compared to the wild type (Fig. 8).
4.4.1 Mechanisms Involved in Rhizobacteria Induced Systemic Resistance
There appears to be lots of variations in terms of the mechanisms underlying the

rhizobacteria induced systemic resistance depending upon the bacterial strain and the
plant involved (van loon 2007). However the presence of reorganization factors
specific to the host plant and the rhizobacterial strain is a must (Pieterse et al. 2002).
The signaling components involved are largely unknown and various kinds of biotic
and abiotic agents may acts as inducers of induced systemic resistance. Biotic agents
include pathogens, non-pathogens and elicitors of fungal cell wall metabolites while
abiotic agents are salicylic acid, ethylene, dichloro-isonicotinic acid and benzothiadi-
azole (Ramamoorthy et al. 2001). Two contrasting set of views have been observed in
Induced Resistance
in Plants
Rhizobacteria
Pathogen
Induced Systemic Resistance (ISR) Systemic acquired resistance (SAR)
Role of SA SA and
and PR proteins Tissue
No visible
PR proteins in always Necrosis
symptoms
elicidation is Enhanced defensive involved in
variable capacity elicidation
Fig. 8 Induced resistance in plants. Though both kinds of induced resistance are phenotypically
similar, inducing agent and elicidaion mechanism are different in both cases. However both result
in activation of latent resistant mechanisms and hence elevated levels of defensive capacity.
SA-Salicylic acid; PR-Pathogenesis related
the literature. As reviewed by Whipps 2001; Pieterse et al. 2002 and supported by
many studies (Hoffland et al. 1995; van Wees et al. 1997; Pieterse et al. 1998), in case
of induced systemic resistance, the induction does not involve pathogenesis–related
proteins like chitinases, b-1, 3 glucanases, protease inhibitors and salicylic acid, it
rather involves jasmonic acid and ethylene mediated signaling pathway that trigger
the defense reaction. Natriuretic peptide receptor A, also known as NPR1 has also
been observed as a regulatory factor for induced systemic resistance (Pieterse et al.
1998; Bargabus-Larson and Jacobsen (2007). However, systemic resistance elicida-
tion in many plants has been observed to be accompanied by an increase in activities
of enzymes like chitinase, ß-glucanase (Bargabus-Larson et al. 2002) peroxidase,
polyphenol oxidase, and phenylalanine ammonia lyase (Chen et al. 2000; Magnin-
Robert et al. 2007), and induced expression of stress-related genes (Verhagen et al.
2004) as well as induction of specific pathogenesis–related proteins in leaves of plants
of which the roots were colonized by resistance-inducing PGPR (van Loon 2007).
The rhizobacterial molecules that have been characterized as determinants of
systemic resistance in plants include lipopolysaccharides (van Peer and Schippers
1992; Leeman et al. 1995b; van Wees et al. 1997), various kinds of siderophores like
pyoverdine-type in Pseudomonas flourecens CHA0 against tobacco necrosis virus
of tobacco, catechol type in Serratia marcescens against Colletotrichum orbiculare
in cucumber, pyocyanin and pyochelin type in Pseudomonas aeruginosa 7NSK2
against Botrytis cinerea in tomato (Maurhofer et al. 1994; Audenaert et al. 2002;
Press et al. 2001) flagella (Meziane et al. 2005) and certain antibiotics like pyocya-
nin, 2,4-diacetylphloroglucinol by P. fluorescens Q2-87 in grapevine against Botrytis
cinerea (Verhagen et al. 2010). Volatile organic compounds produced by certain
Bacillus strains have also been shown as initiators of defense responses in plants.
For example low-molecular weight hydrocarbons including 2,3-butanediol and ace-
toin produced by B. subtilis GB03 and B. amyloliquefaciens IN937a were shown to
induce resistance in Arabidopsis against Erwinia carotovora (Ryu et al. 2004).
Salicylic acid is an iron-chelating molecule with comparatively weaker sidero-
phoric activities. However, the role of salicylic acid accumulation has come out as a
major point of distinction between induced systemic resistance and systemic
acquired resistance. In case of systemic acquired resistance, its elicitation is always
accompanied by accumulation of salicylic acid and plant pathogenesis-related pro-
teins and H2O2 may act as a signaling agent of salicylic acid and an inducer for
activation of PR-I gene (Ryals et al. 1996). Contrasting reports have been obtained
for the role of salicylic acid in elicitation of induced systemic resistance. Several
studies have reported a salicylic acid dependent rhizobacteria elicidated induced
systemic resistance. For example P. aeruginosa 7NSK2 has been shown to elicit
induced systemic resistance in salicylic acid dependent mode (de Meyer and Höfte
1997; de Meyer et al. 1999a). Using mutants of this strain with altered salicylic acid
production abilities, de Meyer and Höfte (1997) demonstrated that induction of
resistance to Botrytis cinerea by P. aeruginosa 7NSK2 was iron-regulated and sali-
cylic acid production is essentially required. Amongst the other two siderphores
produced by this strain i.e. pyochelin and pyoverdin, role of pyochelin was also
established. Similarly de Meyer et al. (1999a) confirmed the role of salicylic acid to
induce systemic resistance in tobacco with Pseudomonas aeruginosa 7NSK2. They
also observed that a pathogenesis-related protein, PR1 was not involved in this
mechanism. At the same time, several other authors have given contrasting reports.
Audenaert et al. (2002) analyzed the role salicylic acid in induction of resis-
tance against Botrytis cinerea in tomato by siderophore producing Pseudomonas
aeruginosa 7NSK2. They used a mutant strain KMPCH (pyochelin -negative,
salicylic acid positive) and showed that pyocyanin and pyochelin, rather than salicylic
acid, contributed to the induced resistance. However, an increase in free sali-
cylic acid on tomato roots colonized by mutant KMPCH (pyochelin negative,
salicylic acid positive) was observed due to in planta salicylic acid biosynthesis.
Similar observation was made by de Meyer et al. (1999b) in bean plants inoculated
with P. aeruginosa KMPCH. Investigations made by Verhagen et al. (2010) with the
same strain of P. aeruginosa i.e. 7NSK2 and its mutants KMPCH (pyochelin and
pyoverdin negative) and KMPCH-567 (pyochelin, pyoverdin, and salicylic acid
negative) established the role of salicylic acid, pyochelin, and/or pyoverdin in prim-
ing phytoalexin responses and induction of resistance in grapevine by 7NSK2
against Botrytis cinerea. Using salicylic acid non accumulating transgenic NahG
Arabidopsis plants Ran et al. (2005) examined salicylic acid producing P. fluorescens
strains WCS374r WCS417r, CHA0r, and 7NSK2 to induce systemic resistance in
A. thaliana against bacterial speck, caused by P. syringae pv. tomato. It was observed
that for all the strains salicylic acid is not involved in elicidation of induced systemic
resistance and some other bacterial determinants have the role to play. Another study
reported the PGPR strains Serratia marcescens 90–166, Bacillus pumilus SE34 and
Pseudomonas fluorescens 89B-61 that induced systemic resistance against blue mold
disease of tobacco caused by Peronospora tabacina were found to operate in a salicylic

acid independent pathway which was not associated with pathogenesis related proteins
(Zhang et al. 2002). Similarly Bargabus-Larson and Jacobsen (2007) described a sali-
cylic acid independent induced systemic resistance elicitation in sugar beet by Bacillus
mycoides isolate BmJ against Cercospora leaf spot. BmJ treatment was followed by
constant level of total salicylic acid in sugarbeet and natriuretic peptide receptor A
activation. This isolate was earlier described to elicit chitinase, ß-glucanase (Bargabus-
Larson et al. 2002) and hydrogen peroxide (Bargabus-Larson et al. 2003) production in
sugar beet which are the biochemical changes normally associated with salicylic acid
dependent signaling pathway. Hence, it was concluded that BmJ activates a salicylic
acid dependent signaling cascade downstream of salicylic acid and systemic resistance
is activated by a novel signaling compound. In their study, using two-strain mixture of
B.subtilis GB03 and B. amyloliquefaciens IN937, Ryu et al. (2007) reported that this
biopreparation protected Arabidopsis against cucumber mosaic virus by a salicylic acid
independent signaling pathway, whereas the protection elicited by the same strain mix-
ture against P. syringae pv. tomato in tomato plants was salicylic acid dependent.
The role of pathogenesis related proteins in induced systemic resistance elicida-
tion in plants like radish has also been ruled out. Various reports support the
redundancy between salicylic acid, jasmonic acid and ethylene signaling pathways.
For example Ryu et al. (2004) reported variable dependence of volatile organic
compounds induced systemic resistance of two stains of Bacillus in Arabidopsis
plants. Strain IN937a triggered induced systemic resistance through an ethylene-
independent signaling pathway, whereas volatile organic compounds from strain
GB03 operated through an ethylene-dependent pathway in triggering induced
systemic resistance. Similarly Ran et al. (2005) reported ethylene dependent induced
systemic resistance by Pseudomonas fluorescens strains WCS374r in Arabidopsis.
Thus, it may be said that induced systemic resistance elicitation may be accompa-
nied by an increase in free salicylic acid in the rhizosphere, but it does not always
act as the signaling compound for the same and more than one kind of mechanism
may be employed by rhizobacteria mediated induced systemic résistance. Other
molecules like jasmonic acid and ethylene also qualify for the class of signaling
components for induced systemic resistance but their exact role remains ambiguous
and may vary with the plant and rhizobacteria species.
5 Conclusion
Various strains of rhizobacteria, Pseudomonas and Bacillus, more particularly, have

come out as promising options to control soil-borne plant pathogens. They have given
satisfactory results under greenhouse conditions and reasonably acceptable perfor-
mance has been observed under field conditions. The range of target pathogens and
crops for which the results have been obtained is also significantly wide. Most of the
biocontrol rhizobacteria tend to follow more than one kind of approach in controlling
the target pathogen. However, secretion of antibiotics and inducing systemic resistance
in the host plant appears to be most widely followed mechanism by PGPR for patho-
gen inhibition. A rich variety of antibiotics, with varied target range and degree of
effectiveness, are produced by rhizobacteria of various kinds, maximum being secreted
by Pseudomonas. Competition becomes the most important tool of action in cases
where the biocontrol agent and target pathogen share high degree of relative proximity
because of the similar requirements in terms of nutrients and growth factors. Role of
siderphores is also irreplaceable as they contribute in many ways ranging from indi-
rect disease control by promoting plant growth, inducing competition among rhizobac-
teria and pathogens to inducing systemic resistance in plants.
However, at present certain issues need attention before moving ahead in this
direction. Two most critical points are economic feasibility of the product and con-
sistency in terms of performance under actual field conditions. Though significant
market growth for microbial/biological tools has been achieved in last few years,
challenge is to attain a competitive level alongside the entrenched and effective
chemical counterparts. Improvement in field performance of these products is the key
to success in this direction. A deeper and unambiguous understanding of the mode
of action of the antagonistic organisms and detailed information about the signaling
molecules involved in the same will help in optimizing their colonization and estab-
lishment in the rhizosphere thereby enhancing the effectiveness.
Further, maximum success so far has been achieved by utilizing two rhizobacte-
rial genera only i.e. Pseudomonas and Bacillus. Therefore, studies for exploring the
vast diversity of rhizospheric microflora must be encouraged in order to widen up
the options available for potential microbial strains for biocontrol. Development of
sensitive and effective molecular tools that may enable the isolation of uncultivable
microorganisms, are highly required to accomplish this task. Hence, it may be con-
cluded that research directed towards the development of large scale productions of
effective, broad range biocontrol agents is the need of the hour. Significantly large
amount of work has already been carried out in this direction, however, to get the
actual commercial success, continuous efforts are required in order to improve the
reliability of these products and to narrow down the gap in terms of product cost of
chemical and biological control tools.
Acknowledgement The authors wish to thank all the persons who helped in design of this manu-
script through their critical comments, valuable suggestions and vital discussions.
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Index
A Alvarez, C.E., 270, 275

Abiven, S., 190 Amensalism, 137–138
Actual yield lose/gain (AYL) index, 131 Amlinger, F., 262
Adebayo, O.S., 324 Andrade, F.H., 106, 110
Afreen, F., 238 Andrews, J., 122, 124
Aggelides, S., 279, 287, 288 Animal breeding
Agriculture, climate changes animal husbandry, 208
African agriculture, 69 CF, 208, 209
agronomic and economic impacts, 69–70 domestic animals, 207
CO2 concentration and enrichment, edible protein, 209, 210
66, 71, 72 food security, 202, 203
crop physiology, 73–76 greenhouse gas, 204–205
cropping systems, 67 higher digestibility, 212–213
crop production, 86–88 higher feed intake, 212
explicit planetary isentropic coordinate livestock revolution, 208
model, 66 lower maintenance requirements, 213
food and forestry trades, 72 lower methane emissions, 213–214
food supply vulnerability, 66 low-phytate maize and pigs, 217
grain yields, 73 nutrient precursor (ß-carotene), 214–216
net primary productivity, 66 nutritionists, 217
O3 concentration, 71, 72 policy and agricultural research, 202, 203
perennial crops, 70 stearidonic soybeans, 216–217
River Nile (see Nile) strategies, 210
simulation models, 70 Anitha, A., 324
temperature and water regimes, 66 Annabi, M., 279
water resources, 70–71 Antibiosis
water scarcity, 76–78 Agrobacterium, 325
water stress, 72 Bacillus species, 325, 329
yield reductions, 73 genetic systems, 326
Agrobacterium, 325 metabolic agents, 323
Agro-ecosystems. See Maize phenazine, 326
Agroforestry, 120–121 Pseudomonas, 325, 327–328
Albaladejo, J., 284 role of, 325
Albiach, R., 291 Serratia, 325, 331
Alexandria, 85 soil microorganisms, 323
Altieri, M.A., 44 Streptomyces, 325, 330
Alva, A.K., 163 systemic resistance, 326
DOI 10.1007/978-94-007-5961-9, © Springer Science+Business Media Dordrecht 2013
362 Index
Apichaisataienchote, B., 330 Serratia, 325, 331

Arabidopsis thaliana, 343 soil microorganisms, 323
Archer, D., 282 Streptomyces, 325, 330
Area harvest equivalent ratio (AHER), systemic resistance, 326
132–133 biopesticides, 322
Area time equivalency ratio and chemical pesticides, 321
(ATER), 132 definition, 322
Aromatic plants. See Medicinal plants ecofriendly agricultural approaches, 323
Arora, N.K., 324 fungal pathogens, 322–324
Arrhenius, S., 43 induced systemic resistance/systemic
Arthur, E., 279–281 acquired resistance
Asaka, O., 329 Arabidopsis thaliana, 343
Audenaert, K., 346 Bacillus spp., 343
Audsley, E., 301, 306 biotic and abiotic agents, 344
Ayyad, M.A., 63 BmJ treatment, 347
KMPCH, 346
lipopolysaccharides, 345
B natriuretic peptide receptor A, 345
Bacillus P. aeruginosa 7NSK2, 346
B. mycoides, 343 pathogenesis, 347
B. subtilis, 323, 324 salicylic acid, 344, 346
B. thuringiensis, 322–324 iron
species Fusarium, 336
antibiosis, 325, 329 P. aeruginosa, 336
induced systemic resistance, 343 pyochelin, 338
Balasubramanian, V., 132 Pythium, 336
Banik, P., 131, 132 siderophores, 336–337
Bargabus-Larson, R.L., 345, 347 microbial inoculants, 322
Bastida, F., 290, 291, 293 parasitism, 340–341
Batlle-Aguilar, J., 187 plant derived nutrients, 338–339
Battu, P.R., 327 root niches
Bauer, W.D., 334 abiotic factors, 336
Bazzoffi, P., 270, 280, 283 agglutination, 334
Beets, W.C., 122, 123 carbon concentrations, 333
Behm, D., 29 cellulose fibrils, 333
Behn, H., 238, 243 chemotaxis, 333
Bell pepper. See N-fertilizer fimbriae, 334
Benbrook, C., 294 gene sss and xerD, 335
Bennett, J., 182 O-antigen lipopolysaccharide, 333
Bernal-Santos, B., 217 P. fluorescens F113, 333
Berzsenyi, Z., 101, 104, 105, 107, P. fluorescens WCS365, 333
109, 114 phenazines, 335
Biala, J., 263 P. putida, 334
Binder, B.Y.K., 237 Pseudomonas, 335
Bingham, G., 33 R. leguminosarum, 333–334
Biocontrol sporulation, 334
antibiosis soil borne phytopathogens, 320
Agrobacterium, 325 starvation, 332
Bacillus species, 325, 329 Biological control agents (BCA).
genetic systems, 326 See Biocontrol
metabolic agents, 323 Biomass, 73
phenazine, 326 Bipfubusa, M., 279
Pseudomonas, 325, 327–328 Bishayee, A., 233
role of, 325 Blair, J.M., 122, 124
Index 363
Bloom, A.J., 169 CO2 concentration and enrichment,

Blumenthal, J.M., 100, 103, 104, 107 66, 71, 72
Böhme, H., 214 crop physiology, 73–76
Boldrin, A., 261, 264, 300 cropping systems, 67
Bonanomi, G., 268 crop production, 86–88
Boomsma, C.R., 106 explicit planetary isentropic coordinate
Brade, W., 211 model, 66
Bradyrhizobium japonicum, 334 food and forestry trades, 72
Bresson, L., 280 food supply vulnerability, 66
Brewer, C., 33 grain yields, 73
Broccoli (Brassica oleracea L.), 169 net primary productivity, 66
Butzer, W.B., 61 Nile River Delta (see Nile River Delta)
O3 concentration, 71, 72
perennial crops, 70
C simulation models, 70
Cameron, R.K., 343 temperature and water regimes, 66
Capdevila, S., 333 water resources, 70–71
Capsicum annuum L. See N-fertilizer water scarcity, 76–78
Carbon dynamics water stress, 72
amino acids, 180–181 yield reductions, 73
bacteria and fungi, role of, 183–184 air temperature, 60
cellulose, 179 annual rainfall, 54
hemicellulose, 179 arid provinces, 63
humification pathways, 181 chemical toxicants, 60
hydrophobic zone, 182 climate disasters, 59
lignin, 179–180 crop yields, 61
modeling of, 184, 186–187 green house gases, 56, 62
organic and organo-mineral compounds, Green Revolution technology, 56
181–182 Hot and dry Khamsin winds, 63
plant-soil system, 184, 185 human activities, 58
Carbon turnover In Pore Space (CIPS) Hyper arid, 63
model, 186 impacts of, 59
Cartwright, D.K., 327 IPCC, 55
Cassava, 126 oceanic process, 55
Castelló, R., 284 population density, 54
Castillejo, J., 284 situation analysis, 63–65
Catharanthus roseus, 240 soil and water resource, 56–58
Cattle, S.R., 25, 27 sub-arctic forests, 59
Celik, I., 274, 288 Colletotrichum orbicularecarnation, 342
Cellulose fibrils, 333 Commensalism, 136–137
Cereals intercrops, 126–128 Competitive ratio (CR), 130–131
Chang, X., 239 Cook, H.F., 276, 282, 283
Charles, R., 306 Coria-Cayupan, Y.S., 274, 294, 295
Chelliah, J., 240 Courtney, R.G., 288
Chet, I., 338, 340 Courty, P.E., 184
Chin-A-Woeng, T.F.C., 324, 327 Cronin, D., 327
Cho, J., 213 Crop physiology
Claassen, V.P., 285 biomass, 73
Clark, J.S., 178 CO2 effects, 75
Climate changes greenhouse gases, 74, 76
agriculture quality of, 75
African agriculture, 69 Crop yield, 269–278
agronomic and economic impacts, Cui, W.H., 233
69–70 Curtis, M.J., 285
364 Index
D Eichholz, I., 239

Daemmgen, U., 211 Ekpo, E.J.A., 324
Davis, J., 301 Elad, Y., 338
DeBach, P., 322 Eldor, P., 182
De Gryze, S., 190, 191 Elherradi, E., 263, 271
Dekkers, L.C., 335 Elsevier, I., 27, 28
de Meyer, G., 346 El Shamy, M.E., 85
De Pascale, S., 294 Enterobacter cloacae, 338–339
de Weert, S., 333 Eriksen, G.N., 270
de Weger, L.A., 333 Everett, P.H., 152
Diacono, M., 262 Explicit planetary isentropic coordinate
Diana, G., 288 (EPIC) model, 66
Diez, J., 270
Dolzhenko, Y., 246
Donna, R., 33 F
Dori, L., 33 Fabrizio, A., 265
Drieling, K., 25–28 Fagnano, M., 266
Drip irrigation system, 145, 147 Falloon, P.D., 186
Duchenne, T., 163 Fanis, L.N., 34
Dungan, K., 33 Farnham, D.E., 100, 102
Dunlop, C.A., 329 Fasoulas, A.C., 110
Duvick, D.N., 110 Fasoula, V.A., 112, 114
Fenglin, H., 230
Fertilizer (Florida)
E crop production, 147–149
Echarte, L., 106, 110 drip irrigation, 145, 147
Education. See Soil education N-fertilizer (see N-fertilizer)
Eghball, B., 265 nitrogen leaching, 144–145, 163, 165
Egypt nitrogen use efficiency, 147–149
climate and climate changes seepage irrigation, 145, 146
agriculture (see Agriculture, climate Fields, S.F., 34
changes) Fiume, F., 324
air temperature, 60 Fiume, G., 324
annual rainfall, 54 Flachowsky, G., 211, 214
arid provinces, 63 Fließbach, A., 291, 293
carbon dioxide, 59 Fortuna, A., 264, 265
chemical toxicants, 60 Francis, C.A., 122, 124
climate disasters, 59 Fridlender, M., 341
crop yields, 61 Frossard, E., 263
green house gases, 56, 62 Full Option Science System (FOSS), 31
Green Revolution technology, 56 Furlough, V., 33
Hot and dry Khamsin winds, 63 Fusarium oxysporum, 340
human activities, 58 Fushimi, S., 330
Hyper arid, 63
impacts of, 59
IPCC, 55 G
oceanic process, 55 García-Gil, J., 291
population density, 54 Gardenas, A.I., 182
situation analysis, 63–65 Geisseler, D., 183
soil and water resource, 56–58 Germ, M., 238
sub-arctic forests, 59 Ge, Y., 292
sustainable agriculture (see Sustainable Geyer, K., 6, 8, 10, 11, 13
agriculture) Ghabbour, S.I., 63
Ehrenfeld, J.G., 184 Ghorbani, R., 274
Index 365
Ghosh, P.K., 132 I

Gibb, L., 33 Iglesias-Jimenez, E., 270, 275
Gil, M., 246 Illera, V., 287
Glab, T., 283 Induced systemic resistance (ISR)
Glandorf, D.C.M., 335 Arabidopsis thaliana, 343
Global circulation model (GFDLT), 85 Bacillus spp., 343
Glucosinolate, 236 biotic and abiotic agents, 344
Golovan, S.P., 213 BmJ treatment, 347
Gómez, E., 292 KMPCH, 346
González, J., 279 lipopolysaccharides, 345
Gosselin, D.C., 19 natriuretic peptide receptor A, 345
Goyal, M.R., 163 P. aeruginosa 7NSK2, 346
Graves, J., 34 pathogenesis, 347
Greenhouse gas emissions (GHG), 208 salicylic acid, 344, 346
Griffiths, B., 290 Inouye, S., 328
Gross National Product (GNP), 81 Intercropping
Guerini, G., 263, 273 actual yield lose/gain index, 131
Gulay, H., 8, 13 aggressivity, 130
Gurusiddaiah, S., 327 agroforestry, 120–121
Gutierrez, V., 157 area harvest equivalent ratio, 132–133
area time equivalency ratio, 132
cereals, 126–128
H competition indices, 128
Hadas, A., 262 competitive ratio, 130–131
Haenel, H.-D., 211 definitions, 122
Hagen, S.F., 235 density
Haggag, W.M., 329 mono cropping yield, 133–134
Haglund, C., 301 mutual cooperation, 134–135
Handoussa, H., 50 mutual inhibition, 134
Hansen, J.C., 33 negative compensation, 136
Hansen, T.L., 263, 300 positive compensation, 135–136
Happs, J.C., 6, 25, 26, 28 predation and parasitism, 135
Hargreaves, J., 262 unilateral cooperation, 136–137
Hargrove, D.L., 33 unilateral inhibition, 137–138
Hartl, W., 276 intercropping advantage index, 132
Hashemi, A.M., 106 land equivalent coefficient, 129
Hassanein, W.A., 328 land equivalent ratio, 128, 129
Hayhoe, D., 20 legume, 126–128
Hemmat, A., 288 Mazaheri’s LER table, 133
High Aswan Dam, 79 mixed intercropping, 122–123
Hillier, J., 304 monetary advantage index, 132
Hochmuth, G., 152 monocultures, 121
Höfte, M., 346 productivity and sustainability, 120
Honeycomb breeding, 110–111, 114 relative crowding coefficient, 130
Hortenstine, C.C., 282, 283 relative yield total, 129
Houck, A., 29 relay intercropping, 124–126
Houot, S., 263 row intercropping, 123–124
Howell, C.R., 327 soil fertility, 121
Hudson, B.D., 282 soil nitrogen, 121
Hulde, M., 9, 16 strip intercropping, 123–125
Human immunodeficiency virus system productivity index, 131
(HIV-1), 240 Intercropping advantage (IA) index, 132
Hwang, B.K., 330 Ioannidis, D., 243, 244
Hypericin, 241 Izaurralde, R.C., 127
366 Index
J crop yield, 269–278

Jackson, L.E., 169 nutrient supply, 261–264
Jacobsen, B.J., 345, 347 soil biological properties and
Jacobsen, E.K., 34 biodiversity, 286, 290–293
Jain, V.C., 230 soil erosion, 278–281
Jamtgard, S., 183 soil moisture content, 278, 282–286
Jastrow, J.D., 180 soil workability, 282, 286–289
Jayabaskaran, C., 237 weed, pest and disease suppression,
Jeremy, C., 33 267–269
Jetiyanon, K., 344 impact assessment methodologies, 260
Johnson, C.B., 239, 240, 243 inventory data, 259
MSW, 257
quantification and impact assessment
K agronomic models, 306
Kamensky, M., 331 carbon sequestration, 301
Karousou, R., 238, 242 crop nutritional quality, 305
Kassam, A.H., 122, 124 crop yield, 302
Kassas, M.A., 47 nutrient supply, 300–301
Keel, C., 327 soil biological properties and
Kennedy, A.C., 33, 34 biodiversity, 304–305
Kilani-Feki, O., 327 soil erosion, 302–303
Kim, B.S., 330 soil moisture content, 303
Kim, P.I., 329 soil workability, 304
Kitchel, T., 29 steady-state models, 306
Kitessa, S.M., 217 weed, pest, and disease suppression,
Kleber, M., 182 301–302
Kloepper, J.W., 344 ReCiPe model, 307
Koch, B., 328 soil loss, 307–308
Kohlrabi (Brassica oleraceagongyloides L.), Lindeijer, E., 307
168 Lin, H.S., 294, 295
Koutroubas, S.D., 98 Lippert, R., 8, 16
Krause, M.S., 343 Litterick, A.M., 267
Kshirsagar, R., 230 Liu, H., 327
Kuka, K., 186 Locascio, S.J., 150, 163
Kumari, R., 237, 238, 242 Londra, P., 279
Loynachan, T.E., 34
Luis, J.C., 239
L Luque, S.F., 110
Laible, G., 214
Lal, R., 264, 304
Land equivalent coefficient (LEC), 129 M
Land equivalent ratio (LER), 128, 129 Macklem-Hurst, J.L., 19
Lap, D.Q., 105 Maffei, M., 242
Lauer, J.G., 100, 102–105 Maize
Laus, M.C., 334 crop management
Lee, C.H., 327 climate conditions, 107
Lee, K.J., 324 drought-prone environments, 107
Legume intercrops, 126–128 early-maturing hybrids, 107
Leroy, B.L.M., 279 high-yielding hybrid, 108
Levenfors, J.J., 331 less population-dependent elite hybrids,
Life cycle assessment (LCA) 109–110
application, 258 population-neutral hybrids, 110–112
compost benefits, 258 short and full-season hybrid, 107–108
carbon sequestration, 264–267 water deficiency, 108
crop nutritional quality, 290, 294–298 modern hybrids, 98–99
Index 367
optimum population variability Monetary advantage index (MAI), 132

environmental variability, 100–102 Montemurro, F., 262, 273, 274
grain yield productivity and stability, Morra, L., 275
102–104 Movahedi Naeini, S.A.R, 276, 282, 283
plant yield potential, 103, 105–106 Mullen, G.J., 288
Malamoud, K., 191, 194 Municipal solid waste (MSW), 257
Mamo, M., 263, 271, 276, 283 Murillo, J., 262
Manzoni, S., 186 Murphy, J.F., 344
Marshall, B., 163 Murray, G.W., 61
Marsh, G.P., 45 Mylavarapu, R.S., 288
Martens, D., 287
Martínez-Blanco, J., 294, 296
Maurhofer, M., 327 N
Medicinal plants Nannipieri, P., 182
aromatic plants Naraghi, L., 324
commercial cultivation, 229 Natriuretic peptide receptor A, 345
global trade, 228–229 Neff, R.L., 34
ayurvedic medicinal plants, 226, 227 Nemecek, T., 304
radical-scavenging and antioxidant Net primary productivity (NPP), 66
potential N-fertilizer
alkaloids, 232–233 agronomic efficiencies, 164
carotenoids, 232 broccoli (Brassica oleracea L.), 169
essential oil, 234 carbon cost, 156
glucosinolates, 234 cost function and management, 151
phenolic compounds, 231–232 crop physiology, 155–156
phytosterols, 233 deeper root systems, 169
polyamines, 234 economic approach, 152
terpenoids, 233 environmental conditions, 151
therapeutic use, 230 irrigation, 168–169
UV-B kohlrabi (Brassica oleraceagongyloides L.),
anthocyanin pigments, 240 168
C. roseus, 240 pepper
flavonoids biosynthesis, 236 crop genotypes, 169
glucosinolate metabolism, 236 daily N uptake rates, 164, 165
hypericin, 241 irrigation, 169
isoprenoids pathway, 245–247 nitrogen uptake efficiency, 148, 166
metabolites enrichments, 247–248 N-rates, effects of, 158–159
phenolics compounds, 236 root dry, 156
polyamines, 241 root uptake efficiency, 169
psoralens, 240 root volume, 169
research, 234–235 soil temperatures, 170
resveratrol, 241 soil zone, 169
shikimate pathway, 245 vegetative and reproductive
volatile oil yield and essential oil, growth, 157
241–244 post-harvest fruit, 153
Mediterranean Sea, 83 potato
Meja, L., 217 daily N uptake, 157, 164, 165
Mildenstein, T., 33 N-rates, effects of, 160–161
Millard, P., 163 root dry, 156
Miller, C.H., 163 root uptake efficiency, 169
Milner, J.L., 329 seepage irrigation, 145, 146
Minasny, B., 187 soil temperatures, 170
Mixed intercropping, 122–123 vegetative and reproductive
Mkhabela, M., 272 growth, 157
Moebius-Clune, B.N., 27, 28 pre-plant, 153
368 Index
N-fertilizer (cont.) Pérez-Piqueres, A., 293

production efficiency, 155–156 Perkins, P., 34
quadratic model, 149, 150 Peronospora tabacina, 347
spinach (Spinaceaoleracea L.), 168 Phenazine, 326
tomato Phillips, D.L., 66
crop genotypes, 169 Pierson, E.A., 327
daily N uptake rates, 164, 165 Pierson, L.S., 327
drip irrigation, 147 Pieterse, C.M.J., 345
fertilizer costs, 151 Piotrowski, J., 33
nitrogen uptake efficiency, 148 Plant breeding
N-rates, effects of, 150, 153, 158 assessment, 217, 218
root growth, 165 climate change, 207
vegetative and reproductive growth, 157 low-phytate maize and pigs, 217
white cabbage (Brassica oleracea L.), 169 nutrient precursor (ß-carotene), 214–216
Ngoundo, M., 303 nutrition, 206
Nielsen, T.H., 327, 328 nutritionists, 217
Niemann, H., 209, 214 objectives, 205–206
Nile River Delta Second Green Revolution, 207
civil society, 79 stearidonic soybeans, 216–217
coastal zone, 82 Plante, A.F., 191
drinking water, 81 Plant growth promoting rhizobacteria (PGPR).
land degradation and desertification, 82 See Biocontrol
rainfall, 80 Popp, M., 101, 107, 108
recycling, 80–81 Porporato, A., 186
sea level, 81, 84 Portnoy, R., 262
temperature variations, 80 Potato. See N-fertilizer
water budget, 80, 82 Pourmorad, F., 230
water demand, in 1997 and 2017, 79 Prakash, D., 230
Nishimura, T., 239 Predation, 135
Nitrogen Pseudomonas
dynamics antibiosis, 325, 327–328
bacteria and fungi, role of, 183–184 root colonization, 335
modeling of, 184, 186–187 Puk, T., 29
in soils, 182–183 Pyochelin siderophore, 338
uptake efficiencies, 166–168 Pyoverdin siderophores, 337
Nitrogen use efficiency (NUE), 147–149 Pythium aphanidermatum, 338–339
Nitz, G.M., 239, 243 Pythium ultimum, 338–339
Norwood, C.A., 101, 107
R
O Rabeeth, M., 324
O-antigen lipopolysaccharide, 333 Rai, R., 237
Odlare, M., 274 Ramani, S., 237, 240
Olness, A., 282 Randler, C., 9, 16
Ouwerkerk, P.B.F., 237 Ran, L.X., 346, 347
Reddy, M.S., 327
Red Sea Hills, 61
P Reinhardt, G.A., 301
Papadopoulos, I.I., 112 Relative yield total (RYT), 129
Parasitism, 135, 340–341 Relay intercropping, 124–126
Particulate organic matter (POM), 188 Rhizobacteria. See Biocontrol
Patel, V.R., 230 Rhizobium
Patyk, A., 301 R. leguminosarum, 333
Pepper. See N-fertilizer R. trifolii, 334
Index 369
Rillig, M.C., 181 Smith, K.L., 34

Root colonization Smith, K.P., 329
abiotic factors, 336 Smith, P., 186
agglutination, 334 Sodhi, G., 265, 278, 279
carbon concentrations, 333 Soil borne phytopathogens, 320
cellulose fibrils, 333 Soil education
chemotaxis, 333 elementary school, 32–34
fimbriae, 334 elementary students
gene sss and xerD, 335 classroom activities, 14
O-antigen lipopolysaccharide, 333 drawings, 11–12
P. fluorescens F113, 333 exploration activities, 10
P. fluorescens WCS365, 333 Gr. 6 British students, 12
phenazines, 335 learning soil layers, 14
P. putida, 334 multiple-choice survey, 10
Pseudomonas, 335 North American school, 11
R. leguminosarum, 333–334 organic and inorganic matter, 15
sporulation, 334 origin, of soil, 15
Ros, M., 273, 291–293 particle size, 15
Rothwell, D.F., 282, 283 post-intervention interviews, 14–15
Row intercropping, 123–124 pre-intervention interviews, 10–11
Russell, T., 9, 10, 14, 16 pre-post gains, 6, 8–9
Rymer, C., 216 pre-post tests, 13, 16, 17
Ryu, C.M., 347 soil comparison, 15
soil nature, 15
soil transformation, 15
S types of soil, 15
Sabrah, R.E.A., 275, 282, 284 urban and rural schools, 11
Said, R., 54 web-based basic soil module, 16
Sánchez-Monedero, M.A., 265 Western European cultures, 11
Sandford, K.S., 61 elementary teachers
Santiago, C.L., 163 environmental survey, 22
Santucci, G., 29 pre-post mean item scores, 24
Sarlangue, T., 105–107 pre-test scores and gains, 23–24
Scannerini, S., 242 pre-test selections, 19–21
Schimel, J.P., 182 soil concepts and attitudes, 22–23
Schnitzler, W.H., 239, 243 equipment kits and resource manuals
Schreiner, M., 239 Full Option Science System, 31
Sclerotium STC unit kits, 30–31
S. rolfsii, 340 teacher resource manuals, 31
S. solani, 340 middle school, 32–34
Seepage irrigation system, 145, 146 secondary school students, 26–27
Sekayange, L., 132 constructivist model, 25
Serratia, 325, 331 hands-on activities, 28
Serratia marcescens, 340 pre-post tests, 28
Severe acute respiratory syndrome secondary school teachers, 29–30, 32–34
(SARS), 240 websites, 32, 35
Shanahan, J.F., 103, 104, 107, 114 Soil erosion, 278–281
Shibu, M.E., 187 Soil moisture content, 278, 282–286
Shoda, M., 329 Soil organic carbon (SOC), 256
Sikora, L.J., 263 Soil organic matter (SOM), 257
Silo-Suh, L.A., 329 carbon dynamics
Simonett, O., 84 amino acids, 180–181
Smith, A., 265 bacteria and fungi, role of, 183–184
Smith, B., 44 cellulose, 179
370 Index
Soil organic matter (SOM) (cont.) definition, 44–45

hemicellulose, 179 Modern Egypt
humification pathways, 181 climate changes, 52
hydrophobic zone, 182 energy sources, 51
lignin, 179–180 food security, 51
modeling of, 184, 186–187 National Urban Development Plan,
organic and organo-mineral 52–54
compounds, 181–182 Nile Valley, 50
plant-soil system, 184, 185 rural poverty, 51
data monitoring, 178 sustainability science, 45–46
nitrogen dynamics sustainable development, 46–47, 50
bacteria and fungi, role of, 183–184 Sutton-Grier, A.E., 290
modeling of, 184, 186–187 Suzuki, S., 284
in soils, 182–183 Systemic acquired resistance. See Induced
soil ecosystem modeling systemic resistance (ISR)
plant feedback mechanisms, 193, 194
principles, 194
state variables, 193 T
soil erosion and degradation leads, 176, 177 Tapia, M.L., 157
structure modeling Taylor, A., 33
empirical models, 190 Taylor, C., 34
limitations of, 191–192 Tejada, M., 276, 279, 280, 287, 290, 291
macroaggregate formation, 188, 190 Terman, G., 262
mechanistic models, 191 Thaning, C., 331
microaggregate formation, 188 Thomashow, L.S., 327, 328
POM, 188 Thomison, P.R., 105, 109, 114
soil aggregate models, 191 Thoppil, R.J., 233
soil stability and fertility models, Thorup-Kristensen, K., 169
188, 189 Tokatlidis, I.S., 98, 101, 104–107, 109, 111,
terrace maintenance, lack of, 176, 177 112, 114
terrestrial ecosystems, 178–179 Tomato. See N-fertilizer
tilling and agricultural practices, 176 Thevenot, M., 180
Speeding, C.R.W., 44 Trichoderma, 340
Spinach (Spinaceaoleracea L.), 168 Trichoderma hamatum 382, 343
Stamatiadis, S., 287 Tuma, J., 238
Stanger, T.F., 100, 102–105 Tumova, L., 238
Stearidonic acid (SDA), 216–217 Turner, M., 287
Stenotrophomonas maltophilia, 341
Sterling, D.R., 33
Stevenson, J.F., 181 U
Stinner, B.R., 122, 124 Ubi, B.E., 240
Stipanovic, R.D., 327 Ultraviolet-B (UV-B)
Streptomyces, 323–325, 330 metabolites enrichments, 247–248
Strip intercropping, 123–125 research, 234–235
Strzepek, K.M., 85 secondary metabolic pathways
Stubbs, T.L., 33 isoprenoids pathway, 245–247
Sullivan, D., 263 shikimate pathway, 245
Sun, M., 238 stress
Sustainable agriculture anthocyanin pigments, 240
Ancient Egypt C. roseus, 240
environmental policy, 48 flavonoids biosynthesis, 236
farming, 49 glucosinolate metabolism, 236
flooding season, 49–50 hypericin, 241
food supply, 48–49 phenolics compounds, 236
Index 371
polyamines, 241 Weller, D.M., 328

psoralens, 240 Whipps, J.M., 345
resveratrol, 241 White cabbage (Brassica oleracea L.), 169
volatile oil yield and essential oil, 241–244 Willey, R.W., 122, 123
United Nations Framework Convention on Wind, B.D., 300
Climate Change (UNFCCC), 64 Wingenbach, G.J., 29
Upadhyay, S., 230 Wolkowski, R.P., 271
Wynen, E., 263
V
van den Boogaard, R., 169–170 X
Vandermeer, J.H., 122, 123 Xanthomonas campestris pv.
Van Meter, D.E., 29 armoraciae, 343
van Peer, R., 342
Verhagen, B.W., 346
Veronesi, P., 33 Y
Vesper, S.J., 334 Yan, W., 99, 110
Vincent, M.N., 327 Yates, D.N., 85
Yoshida, S., 329
Young, P., 217
W Yunlong, C., 44
Wallace, D.H., 99, 110
Wallender, W.W., 300
Wang, J., 329 Z
Wang, S.Y., 294, 295 Zakaria, Z.A., 230
Wang, Y., 236 Zak, J., 2
Wardle, D.A., 180, 184 Zebarth, B., 287
Warman, P., 272 Zehetmeier, M., 212
Watson, S.B., 33 Zheng, X.Q., 237
Weber, J., 282, 285, 289 Zinati, G.M., 288
Weidema, B.P., 307 Zotarelli, L., 164
Wei, G., 342 Zu, Y.-G., 239, 240

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Sustainable Agriculture Reviews

For further volumes:

Call for review articles

ISSN 2210-4410 ISSN 2210-4429 (electronic)

Library of Congress Control Number: 2013932220

© Springer Science+Business Media Dordrecht 2013

Printed on acid-free paper

Springer is part of Springer Science+Business Media (www.springer.com)

Surprising Facts About Soils, Students and Teachers!

Assessing the Environmental Benefits of Compost

Index ................................................................................................................. 361

E. Lichtfouse (ed.), Sustainable Agriculture Reviews, Sustainable Agriculture Reviews 12, 1

the three-dimensional nature of soil, as well as start to understand its formation

Keywords Soil concepts • Sustainability • Education • Hands-on activities • Initial

Science education research – students’ initial understandings of science concepts,

Texas (TEA 2010) In Gr. 1, students sort components of soil by size,

Collins documents a declining trend of the number of undergraduate students

2 Results of Research on Elementary and Secondary

2.1 Elementary Students’ Understanding of Soil

2.1.1 Initial Understandings

changes to clay and then to rock

Table 3 Pre-post gains in soil understanding by elementary students in various countries

intervention their reasons owes its origin to

Fig. 2 A mind-map of children’s previous concepts of soil (Geyer et al. 2004)

2.1.2 The Effect of Treatment Activities

Five studies have attempted to quantify the effectiveness of various “treatment” or

2.2 Elementary Teachers’ Understanding of Soils

Elementary teachers’ understanding about soil may be just as important as that of

2.2.1 Initial Understandings

2.2.2 The Effect of Treatment Activities

In a follow-up study by Hayhoe et al. (Manuscript submitted for publication), 19

… none of the above 30

Fig. 6 Teacher 75% 77%

Fig. 7 Pre-post gains in 88%

Table 8 Pre-test scores and gains on 25 items related to specific soil

2.3 Secondary School Students’ Understanding of Soils

Only two studies report on secondary school students’ initial understandings of

Table 10 Initial understandings of soil by secondary students in three countries

tion of soil components and

2.4 Secondary School Teachers’ Knowledge of and Comfort

There are no published studies I am aware of concerning initial understanding of

3 Resources Available for Elementary and Secondary

3.1 Equipment Kits and Resource Manuals

3.1.1 Complete Equipment Kits

3.1.2 Teacher Resource Manuals

3.2 Short Articles on Soil for Elementary, Middle School,

3.3 Websites and Other Electronic Resources for Soil Education

Grade level Title Description Journal Author(s) Date

of soils” properties using a minimum

Table 13 Websites with soil education resources

Hassan R. El-Ramady, Samia M. El-Marsafawy, and Lowell N. Lewis

H.R. El-Ramady (*)

E. Lichtfouse (ed.), Sustainable Agriculture Reviews, Sustainable Agriculture Reviews 12, 41

Keywords Sustainability science • Climate changes • Sustainable agriculture

2 Sustainable Agriculture in Egypt

Some researchers define sustainable agriculture primarily as a technical process.

2.1 Definition of Agriculture

Therefore, agriculture also called farming or husbandry is the cultivation of ani-

2.2 Sustainability Science

Sustainability is the core element of government policies, university research proj-

The Sustainability Equation

Human Progress & Quality of Life ≤ Ecosystem Goods & Services

Therefore, sustainability science has emerged in the twenty-first century as a new