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Cover Photo and Concept by Dr Pablo Calviño. Commercial-scale test of
sunflower-soybean intercropping in the Pampas of Argentina. Originally a ”subsistence
farming“ concept, here intercropping is a high-tech solution in one of the world’s more
technologically advanced agricultural systems. Crop physiology and enabling agronomy
are behind this innovative technology, as explained in Chapter 3, Section 5.
Academic Press is an imprint of Elsevier
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09 10 11 10 9 8 7 6 5 4 3 2 1
Preface
The problem of agriculture is that it has been too successful
Elias Fereres

As a consequence of the success of post-World War II agriculture, particularly in western Europe, many in
affluent societies have taken food for granted for decades. We lack historical perspective to conclude that the
gap between the world demand and supply of food is widening or otherwise. Nonetheless, the recent debate
on this gap and the role of research and development in agriculture is a positive signal. Nature (2008) has
editorialised on this topic, highlighting the need to spend more on agricultural science to overcome food
crises, whereas the point has also been made that not only the amount but also the allocation of research
efforts is important (Struik et al., 2007).
Current research efforts seem to be under divergent selection favouring either the very large or the very small.
On the one hand, legitimate environmental concerns stimulate investments on global-scale issues. On the
other hand, the internal dynamics of sciences at the molecular end of the scale, where progress is made at
astonishing rates, has become a strong attractor of resources. Crop physiology belongs to the middle ground
between these extremes.
There are many hierarchical levels of biological organisation, from molecules to ecosystems. When we search
for an understanding of biological phenomena, it is commonly found at levels below that of occurrence.
Agro-ecosystem events are explained at the level of the crop, while molecular and cell biology will provide
explanations to physiological responses. Besides, crop physiology provides a vital link between molecular
biology and the agro-ecosystem.
The peak of crop physiology appears to be in the past. Membership in the Crop Physiology and Metabolism
Division of the Crop Science Society of America has declined concurrently with the initiation and rise of the
Genomics, Molecular Genetics, and Biotechnology Division (Boote and Sinclair, 2006). This is a worldwide, rather
than local, phenomenon, and a clear reflection of the shifts in research perspectives towards, in this case, the small.
The objective of this book is to provide a contemporary appreciation of crop physiology as a mature sci-
entific discipline. We want to show that much unfinished business lies in the domain of crop physiology,
and that this intellectually challenging discipline is relevant to agriculture. Progress in agriculture, however,
depends directly on progress in agronomy, plant breeding and their interaction. Hence crop physiology can
contribute to agriculture only to the extent that it is meaningfully engaged with breeding and agronomy;
this is the theme of this book.
V.O. Sadras, Adelaide
D.F. Calderni, Valdivia
© 2009,
2009 Elsevier Inc. xiii
xiv Preface

Boote, K.J. and Sinclair, T.R., 2006. Crop physiology: Significant discoveries and our changing perspective on
research. Crop Sci. 46, 2270–2277.
Nature, 2008. A research menu. Nature 453, 1–2.
Struik, P.C., Cassman, K.G., Koorneef, M., 2007. A dialogue on interdisciplinary collaboration to bridge
the gap between plant genomics and crop science. In: J.H.J. Spiertz, P.C. Struik and H.H. Van Laar (Eds.),
Scale and Complexity in Plant Systems Research: Gene-Plant-Crop Relations. Springer, Dordrecht,
The Netherlands, pp. 319–328.
Contributors

Luis Aguirrezábal INTA-Universidad de Mar del Plata and CONICET, Balcarce, Argentina�
Vincent Allard Institut National de la Recherche Agronomique (INRA), Clermont-Ferrand, France
Fernando H. Andrade INTA-Universidad de Mar del Plata and CONICET, Balcarce, Argentina�
Maria Appendino Universidad de Buenos Aires, Buenos Aires, Argentina
Senthold Asseng CSIRO Division of Plant Industry, Wembley, Australia
Michael Ayliffe CSIRO Division of Plant Industry, Canberra, Australia
Robert Belford Curtin University of Technology, Northam, Australia
Grazia M. Borrelli CRA – Centre for Cereal Research of Foggia, Foggia, Italy
Jeremy J. Burdon CSIRO Division of Plant Industry, Canberra, Australia
Daniel Calderini Universidad Austral de Chile, Valdivia, Chile
Pablo Calviño AACREA, El Tejar, Saladillo, Argentina
Weixing Cao Nanjing Agricultural University, Nanjiing, China
Luigi Cattivelli CRA – Centre for Cereal Research of Foggia, Foggia, Italy
David Connor University of Melbourne, Melbourne, Australia
Mark Cooper DuPont Agriculture & Nutrition, Johnston, IA, USA
Pasquale De Vita CRA – Centre for Cereal Research of Foggia, Foggia, Italy
R. Ford Denison University of Minnesota, Minnesota, MN, USA
Zhanshan Dong DuPont Agriculture & Nutrition, Johnston, IA, USA
Elias Fereres IAS-CSIC and University of Cordoba, Córdoba, Spain
Ralph A. Fischer CSIRO Division of Plant Industry, Canberra, Australia
M. John Foulkes University of Nottingham, Leicestershire, UK
François Gastal Institut National de la Recherche Agronomique (INRA), Lusignan, France
Gurjeet S. Gill University of Adelaide; School of Agriculture, Food & Wine, Waite Campus, Australia
Victoria Gonzalez-Dugo University of Córdoba, Córdoba, ������
Spain�

© 2009,
2009 Elsevier Inc. xv
xvi Contributors

David Guest University of Sydney, Sydney, Australia

Kaija Hakala MTT Agrifood Research Finland, Jokioinen, Finland
Antonio Hall Universidad de Buenos Aires-IFEVA and CONICET, Buenos Aires, Argentina
Graeme Hammer University of Queensland, Brisbane, Australia
Natalia Izquierdo INTA-Universidad de Mar del Plata and CONICET, Balcarce, Argentina�
Hannu Känkänen MTT Agrifood Research Finland, Jokioinen, Finland
Adriana C. Kantolic Universidad de Buenos Aires, Buenos Aires, Argentina�
Armen R. Kemanian Texas Agricultural Experiment Station, Georgetown, TX, USA
Gilles Lemaire Institut National de la Recherche Agronomique (INRA), Lusignan, France
Fulco Ludwig Earth Systems Science & Climate Change Group, Wageningen University, Wageningen, The
Netherlands
Pierre Martre Institut National de la Recherche Agronomique (INRA), Clermont-Ferrand, France
Anna M. Mastrangelo CRA – Centre for Cereal Research of Foggia, Foggia, Italy
Glenn K. McDonald University of Adelaide; School of Agriculture, Food & Wine, Waite Campus, Australia�
Carlos Messina DuPont Agriculture & Nutrition, Johnston, IA, USA
Daniel Miralles Universidad de Buenos Aires-IFEVA and CONICET, Buenos Aires, Argentina�
Juan P. Monzon AACREA, UNMdP, El Tejar, Saladillo, Argentina
Jairo Palta CSIRO Division of Plant Industry, Wembley, Australia
Robert F. Park University of Sydney, Sydney, Australia
Pirjo Peltonen-Sainio MTT Agrifood Research Finland, Jokioinen, Finland
Gustavo Pereyra-Irujo INTA-Universidad de Mar del Plata and CONICET, Balcarce, Argentina�
Dean Podlich DuPont Agriculture & Nutrition, Johnston, IA, USA
Ana C. Pontaroli INTA-Universidad de Mar del Plata, Balcarce, Argentina
Ari Rajala MTT Agrifood Research Finland, Jokioinen, Finland
Matthew P. Reynolds CIMMYT, DF, Mexico
Andrew J. Robson Queensland Department of Primary Industries and Fisheries, Queensland, Australia
Daniel Rodriguez Queensland Department of Primary Industries and Fisheries, Queensland, Australia
Victor Sadras South Australian Research & Development Institute, Adelaide, Australia
Rodrigo G. Sala INTA-Universidad de Mar del Plata, Balcarce, Argentina�
Roxana Savin University of Lleida, Lleida, Spain
Ram C. Sharma CIMMYT, Kathmandu, Nepal
Gustavo A. Slafer ICREA & University of Lleida, Lleida, Spain
 Contributors xvii

Claudio O. Stöckle Washington State University, Pullman, WA, USA

Roger Sylvester-Bradley ADAS, Cambridge, UK
Michelle Watt CSIRO Division of Plant Industry, Canberra, Australia
Weijian Zhang Nanjing Agricultural University, Nanjiing, China
Acknowledgements

Professor Antonio J. Hall contributed to maintain the focus of the book in difficult times, and provided
insightful perspectives for many of the chapters in this book. Only his modesty prevented this book to count
him as third editor.
We thank the contribution of all the authors; their knowledge and time made this book possible. Reviewers
who provided valuable input to one or more chapters include John Angus, Ken Boote, David Connor, Julio
Dardanelli, Jenny Davidson, Tony Fischer, Antonio J. Hall, Anthony Hall, Peter Hayman, John Kirkegaard,
Will Ratcliff and Huub Spiertz.
We are grateful to our host organisations, the South Australian Research and Development Institute and
Universidad Austral de Chile, for their support to this project.
We thank Elsevier’s staff for supporting this project, especially Ms. Pat Gonzalez, Mr Mani Prabakaran and
Dr. Andy Richford for their professional advise and support, and Ms. Christine Minihane for her work at
early stages of the book.
Throughout the book, key concepts developed by the authors are supported with material previously pub-
lished in several journals. We thank the publishers who permitted to reproduce their material: American
Society of Agronomy, Inc. (ASA), Crop Science Society of America, Inc. (CSSA) and Soil Science Society
of America, Inc. (SSSA), American Society of Civil Engineers Publications (ASCE Publications), CSIRO
Publishing, Édition Diffusion Presse Sciences (EDP Sciences), Elsevier, Food Product Press, Italian Society
of Agronomy, Kluwer Academic Publisher, Limagrain, Oxford University Press, Springer, The Haworth Press,
Weed Science Society of America and Wiley-Blackwell Publishers.
Ana Ruben and Magda Lobnik helped with editorial checks and supported the editors with loving patience.

2009 Elsevier Inc.

© 2009, xix
 CHAPTER 1

Sustainable Agriculture and Crop

Physiology
Victor Sadras, Daniel Calderini and David Connor

… the herd of cattle was an animal who wanted to be many, or many animals that wanted
to be one…
Ricardo Güiraldes

1. Introduction
A view from space emphasises the areal dimension of agriculture (Loomis and Connor, 1996), which is
threefold. First, the spreading of crops across the landscape allows for the effective capture of both aerial
(sunlight, CO2) and soil (water, nutrients) resources that sustain crop growth (Loomis and Connor, 1996;
Monteith, 1994). The linkage between spatial distribution of resources and plant growth requirements leads
to predictable large-scale patterns of spatial distribution of vegetation (Miller et al., 2007), and competition
for resources influences the structure and functioning of plant populations and communities (Benjamin and
Park, 2007; Donald, 1981; Dybzinski and Tilman, 2007; Harper, 1977; Tilman, 1990). Second, smaller-scale
space-dependent plant-to-plant interactions mediated by a wide array of signals allow plants to differenti-
ate self and no-self, and respond to their neighbours before competition for resources arises (Aphalo and
Ballaré, 1995; Aphalo and Rikala, 2006; Ballaré et al., 1990; Djakovic-Petrovic et al., 2007; Dudley and File,
2007; Karban, 2008; McConnaughay and Bazzaz, 1991). Third, there is an areal dimension in the defini-
tion of communities and ecosystems where the most significant aspects of agriculture are defined, including
yield (Box 1) and resource conservation. A typical cropping field includes a thropic web of primary pro-
ducers (crop species and weeds), and primary and secondary consumers, including herbivores, pathogens
and predators; decomposers play a critical role in the geochemical cycles of major nutrients. Collectively,
all these organisms constitute a community, which together with the physical and chemical attributes of the
environment form an agro-ecosystem (Loomis and Connor, 1996).
Crop physiology deals with the structure and functioning of crops, and is therefore closely related to plant
sciences and ecology. The principles of crop physiology have been reviewed comprehensively, and are there-
fore out of the scope of this book. Readers interested in these principles are referred to a series of classi-
cal and more recent books (Charles-Edwards, 1982; Evans, 1975, 1993; Fageira et al., 2006; Gardener
et al., 1985; Hay and Porter, 2006; Johnson, 1981; Loomis and Connor, 1996; Milthorpe and Moorby, 1979;
Pessarakli, 1995; Smith and Hamel, 1999).
In this chapter, we outline the paradigm of contemporary agriculture, and the challenge of sustainable pro-
duction of bulk and quality food, fodder, fibre and energy during the next decades (Sections 2 and 3). We
assess critically a range of agricultural aims and practices, including organic farming and production of
land-based raw materials for biofuels; the environmental, economic or social flaws of these approaches are

2009 Elsevier Inc.

© 2009, 
 CHAPTER 1: Sustainable Agriculture and Crop Physiology

BOX 1  Crop yield: An evolving concept with multiple definitions

Evolution of yield criteria (e.g. Cassman et al., 2003) could be a consequence of
For birds and mammals, natural selection favours the increasing cropping intensity. This is nicely illustrated by
procurement of food at the lowest energy cost within the Egli (2008), who reported an inverse relationship between
phylogenetic constraints of neural architecture (McLean, the rate of progress of the yield of soybean crops and the
2001). Before agriculture, our ancestors were therefore intensity of cropping measured as proportion of double
not unlike other animals, for which ‘yield’ was the ratio crops in the system: the relative rate of yield improve-
between the energy derived from food and the energy ment from 1972 to 2003 declined from 1.51% in counties
invested in obtaining food (Schülke et al., 2006). In the with little or no double cropping to 0.66% in counties with
transition from foraging to farming, human populations 70% of land allocated to wheat-soybean double cropping.
were exposed to strong selective pressures to capture the Paradoxically, the best environments supporting higher
full benefit of an energy-dense but initially detrimental cropping intensity could therefore show the lower rate
food supply, including starchy cereals, roots and tubers, of improvement in the yield of individual crops. Explicitly
fatty meats, dairy products and alcoholic beverages (Patin measuring yield per unit area and time, as proposed by
and Quintana-Murci, 2008). Although the enzymatic Evans, is therefore of increasing importance.
machinery to metabolise these substances was already in
place, the basal expression of such enzymes in our ances- Yield of individual crops: Definitions
tors was inadequate to process the massive amounts of Yields are in a continuum from crop failure to potential
food brought about by farming. High levels of amylase in (Loomis and Connor, 1996), and several authors have
saliva or lactase in the small intestine probably conferred proposed definitions to account for this range (Bingham,
a strong selective advantage to early farmers (Patin and 1967; Evans and Fischer, 1999; Loomis and Amthor, 1999;
Quintana-Murci, 2008). Loomis and Connor, 1996; van Ittersum and Rabbinge,
Once the sowing of crops was established as a common 1997). In this book, we favour the definitions of actual and
practice, the definition of yield shifted from an energy attainable yield of Loomis and Connor (1996) and the defi-
ratio to the ratio between seed harvested and seed sown nition of yield potential of Evans and Fischer (1999).
(Evans, 1993). This was particularly important in low- n Actual yield is the average yield of a district; it
yielding seasons, when farmers had to make the hard reflects the current state of soils and climate, aver-
decision of allocating seed for food and seed for the next age skills of the farmers, and their average use of
sowing. An important consequence of this measure of technology. Actual yield also applies to the particular
yield was that selection possibly favoured highly com- yield of a crop in a given paddock and season.
petitive plant types, that is, abundant tillering, large inflo-
rescences, small grain and weak seed dormancy (Evans,
n Attainable yield ‘corresponds to the best yield
1993). Only when availability of arable land came under achieved through skilful use of the best available
pressure, mass of product per unit land area became a technology’.
more important criterion. This shift in the definition of n Potential yield is ‘the maximum yield that could be
yield had a dramatic impact on selective pressures, shift- reached by a crop in given environments’ as deter-
ing from the aggressive high-yielding plant (grains per mined from physiological principles. As such, it is
grain) to the less competitive ‘communal plant’ able to distinguished from yield potential, that is, the ‘yield
produce more yield per unit area (Donald, 1981). Evans of a cultivar when grown in environments to which
(1993) envisaged the next measure of yield whereby the it is adapted, with nutrients and water non-limiting
time dimension is considered explicitly, for example, yield and with pests, diseases, weeds, lodging and other
per hectare per year. This measure is particularly impor- stresses effectively controlled’. In this definition, envi-
tant when comparing productivity of systems where the ronment emphasises solar radiation, temperature and
degree of intensification is variable. Intensification of agri- day length (Evans and Fischer, 1999). Building on
culture is a worldwide phenomenon (Caviglia et al., 2004; this definition, we propose that diffuse radiation and
Farahani et al., 1998; Sadras and Roget, 2004; Shaver vapour pressure deficit need explicit consideration,
et al., 2003; Zhang et al., 2008). In this context, some alongside radiation and temperature (Rodriguez and
observations of stabilisation or decline in yield (kg ha1) Sadras, 2007). This is based on known phenomena,

(Continued)
 2. Agricultural Paradigms 

BOX 1  Continued
that is, higher fraction of diffuse radiation favours A ranking of yields is expected:
canopy photosynthesis (Roderick and Farquhar,
Actual yield≤attainable yield
2003; Sinclair and Shiraiwa, 1993; Sinclair et al., 1992),
<potential yield<yield potential
and high vapour pressure deficit restricts growth of
well-watered plants (Bunce, 2006; Gollan et al., 1985; Analysis of gaps between successive yield levels allows
Monteith, 1993; Sadras et al., 1993). for identification of constraints and trade-offs.

emphasised (Section 4). In the context of sustainable agricultural systems, we define the aims and outline
the structure of this book (Section 5).

2. Agricultural paradigms
Agriculture started in the Neolithic, about 10,000–12,000 years ago (Childe, 1927; Flannery, 1973). At the
beginning and for most of its history, the core objective of agriculture was to harness resources into the most
diverse products, from cereal grain and cotton fibre to olive oil and wine. Environmental aspects of crop
production have concerned farmers since the early days of agriculture, and from the mid-nineteenth century,
scientists have also focused on the links between production and resource conservation.
Many farming practices evolved to account for environmental issues that often led to improvements in both
resource protection and production. In early farming systems relying on shifting fallows, one or two seasons
of cropping were often followed by regeneration periods of 20–25 years for forest or 6–10 years for bush
(Evans, 1993). The four-course rotation developed in Norfolk in the eighteenth century comprised seven
elements: enclosures, use of marl and clay, proper rotation of crops, culture of turnips, culture of clover and
ryegrass, long leases, and large farms (Evans, 1993). This early technology illustrates two important aspects
of agricultural innovation. First, it reinforces the notion that farmers are generally aware of the importance
of maintaining their resource base, as shown by the combination of techniques aimed at maintaining soil
fertility. Second, it illustrates the principle that substantial improvements in production generally do not
result from a single innovation, but rather by the ingenious rearrangement of many, often pre-existing tech-
niques and exploitation of synergies (Evans, 1993, 2005).
The contemporary concept of sustainability accounting for the triple bottom line of economic, environmen-
tal and social outcomes has been implicit in agricultural practices and research for a long time, but has only
been formalised in the last four decades. The concept of sustainable agriculture raised its profile exponen-
tially in the 1980s and was firmly established in the 1990s (Figure 1). In tune with this trend, a large num-
ber of long-term agro-ecosystem experiments were established in the 1980s and 1990s.1 Food safety and
more subtle aspects of the nutritional characteristics of food products (Mayer et al., 2008; Trethowan et al.,
2005; Welch and Graham, 1999) are elements of increasing importance that add a fourth dimension to the
contemporary paradigm of agriculture (Figure 2).

1
Rasmussen et al. (1998) defined long-term agro-ecosystem experiments as ‘large-scale field experiments more than 20 years old
that study crop production, nutrient cycling and environmental impacts of agriculture’, and reviewed the features of ‘classic’ (i.e. more
than 50 years old) and more recently established experiments in this class. Examples of classic long-term agro-ecosystem experiments
include those at Rothamsted (established in 1843) in the United Kingdom; Morrow (1876) and Sanborn (1888) in the United States;
Askov (1894) in Denmark; Eternal Rye (1878) and Static Fertiliser (1902) in Germany; Rutherglen (1913), Longerenong (1917) and
Waite (1925) in Australia; Skierniewice (1923) in Poland; and Leeethbridge (1911) and Breton (1930) in Canada. In response to the
renewed importance of questions in long-term time scales, the number of long-term agronomic experiments raised from less than 30
to more than 240 in recent decades (Rasmussen et al., 1998).
 CHAPTER 1: Sustainable Agriculture and Crop Physiology

1000

Number of publications per year

100

10

0.1
1970 1975 1980 1985 1990 1995 2000 2005
Year
Figure 1
Time course of papers retrieved searching for ‘sustainable agriculture’ in the Commonwealth Agricultural Bureaux (CAB) database
(1910–2007). No publications were retrieved for the period 1910–1967. The first paper retrieved with this search criterion
was published in 1968, and its focus was on economic sustainability in relation to land price and alternative land uses, that is,
agricultural versus industrial or urban (Cervinskij et al., 1968). Biswas and Biswas (1976) challenged the sustainability of high-input
agriculture in the context of the energy crisis of the 1970s, and this is one of the first formal accounts, albeit partial, of sustainability
in the contemporary meaning of the concept. Bennett and Schork (1979) explicitly considered the environmental, economic and
social aspects of sustainability, and theirs is the first publication to feature ‘sustainable agriculture’ in the title. The concept of
sustainable agriculture raised its profile exponentially in the 1980s and was firmly established in the 1990s.

Sustainable agriculture

Improves the quality of life of

individuals and communities
value and safety of food
Protects the integrity of

Ensures the nutritional

natural resources

Science
Technology
Policy
Education

Is profitable
Figure 2
Concerted efforts in science, technology, policy and education are required to deliver on the four dimensions of sustainable
agriculture.
 3. World Trends in Population and Demand of Agricultural Products 

The link between sustainability and complexity is an exiting theoretical challenge and a point of huge impli-
cations for policy makers. It could be argued that the environmental dimension of sustainable development
has always been complex, but whether this leads to stasis or collapse is uncertain (Fisk and Kerherve, 2006).
In addition, the growing complexity of social and financial systems has been seen negatively, as complex
systems risk drifting into unstable domains with no means to steer the system out of its course, whereas the
positive side of this problem is a sort of protection against single agents taking control (Fisk and Kerherve,
2006). Finnegan (2003) for instance explained the 2001 outbreak of foot-and-mouth disease in the UK in
terms of complexity. Unlike previous outbreaks, this spread faster and farther: 750,000 cattle and 4.5 million
sheep were slaughtered, a general election was postponed, and a divide was open between country and city.
Finnegan pointed out three elements contributing to the outbreak: (a) economic rationalisation of abattoirs
and bizarre EU subsidies that increased the connections between herds to a critical point,2 (b) changes to
reporting rules that may have delayed the identification and isolation of infectious animals, and (c) fail-
ure to integrate the economics and the biological components of the system; the variables manipulated to
improve economic return also enhanced the rate of spread of the disease.
In the context of sustainability defined along the four axes in Figure 2 and the increasing complexity of
world agricultural systems, the major challenges of agriculture in the next decades relate to the increasing
demands of food and other agricultural products (Section 3), competing land uses of both agricultural and
non-agricultural nature, and misplaced emphasis on ecologically, economically or socially unsound agricul-
tural practices (Section 4).

3. World trends in population and demand of

agricultural products
With justified optimism, it was thought that it was a matter of time to completely solve the problem of food
shortages after the Green Revolution in the later 1960s. This problem was related to poverty and food distribu-
tion as much as it was related to actual production and improved cropping systems. These problems remain for
many reasons, including insufficient and poorly allocated investment in research and development in agriculture
(Anon., 2008; Borrás and Slafer, 2008; Evans, 1997; Marris, 2008; Struik et al., 2007). Furthermore, naive views
of affordable and reliable food supply in affluent societies have been challenged by the convergence of a com-
plex array of factors, including competition for land between food production and alternative uses (Section 4),
climate shifts; increasing price of inputs, particularly fertilisers and energy; and increasing food demand result-
ing from the compounded effect of increasing population and qualitative changes in food demand.
Human population has increased at an impressive pace from 1850 to reach 6 thousand million at the end
of the twentieth century (Evans, 1997; UN, 2007). By the middle of the present century, world population is
estimated to increase to 9.2 thousand million from the current 6.7 thousand million (UN, 2007). The almost
40% increase in population projected for 2050 will be the most powerful driver of food demand; food con-
sumption is predicted to be 3130 kcal person1 day1 in 2050 from 2789 in 2000 (FAO, 2006). Additionally,
qualitative shifts will also influence the demand of agricultural products. In industrialised countries, 73% of
cereals are fed to animals in comparison to 37% in developing countries (Pretty, 2008). Only to satisfy the
increasing demand for animal products in developing countries, an extra 300 Mt of grain will be needed
by 2050 (Pollock et al., 2008). The expansion of agricultural land area was the most important driver of
increased production of main crops during the first half of the twentieth century, and the increase of crop
yield was fundamental afterwards (Miller, 2008; Slafer et al., 1996; Slafer and Otegui, 2000). The challenge
is therefore to further increase production, if anything, with the same or less amount of land and water.

2
In a system with n elements and x random connections between them, the ratio n/x is a measure of integration. The size of the
connected cluster increases slowly with increasing n/x until it reaches a critical point or phase transition, where a giant cluster
‘crystallises’ (Kauffman, 1995). Beyond this point, important properties emerge that cannot be deduced from the parts of the system.
 CHAPTER 1: Sustainable Agriculture and Crop Physiology

Barley Maize Potato

35 90 160
30 80 140
70 120
25
60 100
20 50
80
Grain yield gain (kg ha�1 year�1)

15 40
30 60
10
20 40
5 10 20
0 0 0
Rice Soybean Sunflower
60 35 12
50 30 10
40 25 8
20
30 6
15
20 4
10
10 5 2
0 0 0
1961–2006 1961–1989 1990–2006 1961–2006 1961–1989 1990–2006
Wheat
50

40

30

20

10

0
1961–2006 1961–1989 1990–2006
Period
Figure 3
Global yield gains of barley, maize, potato, rice, soybean, sunflower and wheat calculated for three periods: 1961–2006,
1961–1989 and 1990–2006. Yield gain is the slope of linear regressions between annual global yield and year; bars are the
standard error of the slope.

Figure 3 shows global yield trends of the four more important cereal crops (wheat, rice, maize and barley),
the most widely sown grain legume (soybean) and oilseed crop (sunflower) and the largest non-grain food
crop (potato). Assuming a population of 8.0 thousand million by 2020–2025 and considering that wheat,
rice and maize account for 60% of the human food demand (Tilman et al., 2002), Byrnes and Bumb (1998)
calculated the yields necessary to meet this demand: 5.4 t ha1 for wheat, 5.3 t ha1 for rice and 5.8 t ha1 for
maize. Projecting the yield trends from the last 17 years (Figure 3) into 2020 indicates 3.3 t ha1 for wheat,
4.8 t ha1 for rice and 6.2 t ha1 for maize, representing gaps of 40, 10 and 8%, respectively.
This type of rate analysis has been the standard to detect shifts in productivity trends and gaps between sup-
ply and demand. However, this approach has two drawbacks. First, short-term trends are strongly influenced
by local or global temporary phenomena (e.g. widespread drought or political turmoil in major producing
countries or unfavourable grain-to-fertiliser price ratio). Evans (1997) coined the term ‘plateau spotters’ to
describe the claims that crop yields are approaching their limit on the grounds of short pauses in yield trends,
for example, as seen for wheat and rice in Figure 3. The second reason why yield per unit area and rates
derived from it are inappropriate is that farmers are primarily interested in the productivity and profit of their
farms, rather than in the production of individual crops. In some cases, for example, in the extreme environ-
ments of northern Europe outlined in Chapter 4, yield of individual crops and whole-farm productivity
are tightly related. In most of the temperate to tropical regions of the world, the link is more tenuous owing
 4. Productivity is the Key But Inadequate for all Society’s Demands 

to trade-offs between yield of individual crops and cropping intensity (Box 1). Egli (2008) demonstrated this
strong trade-off for soybeans in the USA in a study analysing county yield trends between 1972 and 2003
(Box 1). Likewise in Argentina and Brazil, double-cropped soybean is increasingly important. Interpretation
of rates of change in yield per unit area of soybean in Figure 3, therefore, needs to account for the trade-
off between soybean yield and whole-farm production and the profit in the three largest world producers
(Figure 1 in Chapter 3). The same applies to many other crops including wheat and rice in both dry land
and irrigated systems. In the central Great Plains of North America, low and unreliable rainfall has favoured
winter wheat-fallow rotations, but greater cropping frequency at the expense of fallow may in some cases
improve farmers’ profitability despite reduced wheat yields (Lyon et al., 2004, 2007). In the more favourable
environment of the Pampas, where continuous cropping has already been established for decades, farmers
are stepping up cropping intensity to improve profitability, often at the expense of yield in individual crops
(Section 5 in Chapter 3). Part of the apparent ‘plateau’ or decline in rice productivity perceived in recent
analyses (e.g. Chapter 9) is related, as already explained by Evans (1997), to the selection for shorter-sea-
son varieties that allow more crops per year: ‘four rice crops grown at Los Baños in the Philippines within
a period of 335 days yielded 76.7 kg grain per hectare per day, giving a total of 25.7 t ha1 for the year, 10%
more than the world record maize crop’. It is clear that tons per hectare is an inappropriate measure of yield
in dynamic systems increasing cropping frequency at rapid rates.
Meeting future world demands of food depend on both improved assemblage of activities (cropping, live-
stock production) in farming systems and further increases in yield of individual crops through the combi-
nation of plant breeding, improved agronomy and their synergy.

4. Productivity is the key but inadequate for

all society’s demands
In a highly populated world, a key to sustainability is large primary production (Section 3). That in turn
requires high productivity of limited available land and can only be achieved by efficient use of germplasm,
labour, managerial skill, energy, water, fertiliser and other agrochemicals. Without high productivity, it is not
possible to provide an adequate and secure food supply for the present or expected world population, while
preserving sufficient non-agricultural land for its other services and values. The magnitude of that challenge
and the role of technology in its achievement were analysed thoroughly in Feeding the Ten Billion (Evans,
1998) and How much land can ten billion people spare for nature (Waggoner, 1994). We do not now expect
a world population of 10 thousand million (10  109), rather 9.2 thousand million by 2050 (UN, 2007);
however, the challenge is more daunting because, on the one hand, supporters of low input and organic
agriculture promote systems that are inherently unproductive, and on the other hand, supporters of bio-
fuel production require enormous increases in productivity for even a small contribution of liquid fuel to
transportation.
Regretfully both movements are widely supported by governments with polemic and subsidies for reasons
that defy logic or science. These challenges to productivity are important because there are questions of food
and environmental safety, energy security and mitigation of climate change. But the question must be asked
if the proposals meet these challenges. Detailed analyses of organic agriculture (Box 2) and land-based bio-
fuels (Box 3) are beyond the scope of this chapter. However, these two industries help to emphasise that a
sustainable future can only be achieved by greater productivity underpinned by improved understanding of
crops and cropping systems, and careful and comprehensive analyses of their attainable yields and environ-
mental impact. Organic agriculture and biofuel are two currently important agricultural issues with signifi-
cant implications for society at large. During the lifetime of this book, some of these issues might fade away.
A long-lasting lesson is, perhaps, to consider these as a property of human nature searching for naive and
easy solutions to complex problems with little regard for their consequences beyond their narrow intended
target (Asimov, 1972). The application of the best science and technology are indispensable elements in the
search for sustainable food production.
 CHAPTER 1: Sustainable Agriculture and Crop Physiology

BOX 2  Misleading comparisons of crop production in organic and

conventional agriculture
The objective of the organic movement is to produce the legumes where nitrogen is supplied by fertiliser. In
safe food in a way that preserves the environment; the regions of high food demand, fertiliser is essential for the
method adopted to ‘ensure’ this is avoidance of synthetic intensive cropping required to provide adequate food.
chemicals and reliance on natural substances for man- Thus in Bangladesh, a highly populated country situated
agement of pests, diseases and fertility. Organic agri- on the once fertile delta of the Brahmaputra and Ganges
culture currently occupies 31 Mha of agricultural land, Rivers, the average cropping intensity is 2.5, that is, two
mostly in developed countries (Willer and Yussefi, 2007). or three crops are harvested from each field each year.
Of that total, 15 Mha found in two countries, Australia In the widespread rice–wheat system there, annual yield
and Argentina, are mostly extensive zero-input grazing of 8–10 t grain ha1 would not be possible in an organic
country. Consequently, the part that is farmed to rules system because, other nutrient issues aside, the fixation
administered by various regulating associations is a small of the required amount of N (approx. 300 kg N ha1) would
proportion (1%) of the 1500 Mha of world cropland. drastically reduce crop area and frequency. In other
Misinterpretation of agronomic principles, including crop places where food supply is not so critically short, inef-
ratios and yield definitions, have led to unsound recom- ficient use of agricultural land in organic systems requires
mendations for organic agriculture, particularly as a solu- more land to meet production requirements, so less land
tion for less developed countries. can be set aside for nature.
Based on crop productivity of ‘organic’ systems of medi-
Crop ratios
eval Europe and assuming all land is devoted to agri-
Crops characteristic of subsistence agriculture, including
culture, Smil (2001, 2004) estimated maximum carrying
beans, maize and cassava, often show yield ratios relative
capacity of organic agriculture in the range of 3–4 thou-
to current management in excess of 2 (Pretty et al., 2003).
sand million, well below the present population. Badgley
Those high ratios are used to promote organic agriculture
et al. (2007) recently sought to refute those estimates, but
as the solution to developing country agriculture (Badgley
their own estimates were badly flawed because they did
et al., 2007). After all, if yield can be doubled without
not account for the severe limitations that the supply of
fertiliser, then why struggle to buy any! The answer is
organic manures has on the expansion of organic agricul-
found in yields achieved rather than ratios. Despite the
ture (Connor, 2008).
increases, the yields remain small. Increasing yield of
maize crops from 0.5 to 1.2 t ha1 (ratio .2), for example,
is hardly a long-term solution to hunger when the same A place for organic farming
gains could be achieved on all fields with small doses of In those places where fertilisers are either not available
fertiliser, and yields of 5–10 t ha1 with larger applications. or too costly and subsistence agriculture predominates,
application of rotations and fertility management prac-
Yield definitions tices can substantially increase the yield of individual
Crops can indeed be grown to equally high yields when crops (Figure 3 in Chapter 5). For these developing coun-
treated with appropriate amounts of inorganic or organic try farmers, organic methods should be fairly presented
fertilisers, but the environmental ‘benefits from reliance as a valuable start to increasing yields and solving hun-
on organic N sources have not been established, and the ger but should not, with their exclusion of fertiliser use,
scientific basis for such a perception has not been docu- be presented as the sustainable solution. Organic agricul-
mented’ (Cassman et al., 2003). Whilst carrying no envi- ture does not adequately address the most urgent needs
ronmental benefit, the task is more difficult with organic of rural poor in developing countries. As in the developed
nutrients because of the large volumes required and the world, it is best implemented as a complementary oppor-
usually small, and mostly unknown, nutrient content. This tunity to generate additional income in peri-urban areas
equality of yield does not, however, establish that organic on land with initial high fertility or where opportunity
agriculture is an equally productive system as alternatives exists to import organic nutrients, and where consum-
using inorganic fertiliser. Other land is required to provide ers are prepared to pay price premiums for the prod-
that organic fertiliser, so the yield of the organic approach ucts. Increasingly, however, organic products are faced
on a system basis, that is, per unit area and time, is with competition from ‘Good Agricultural Practices’ (e.g.
smaller. If a crop can be grown in rotation with legumes EurepGAP) that certify food safety and environmental
in an organic system, additional crops can substitute stewardship using agrochemicals (Hobbs, 2007).
 4. Productivity is the Key But Inadequate for all Society’s Demands 

BOX 3  Land-based biofuels

In response to the oil price shock in the 1970s, agriculture energy costs of transformation to biofuel (chemical trans-
was called to provide liquid fuels, bioethanol from sugar and esterification of vegetable oil to biodiesel is significantly
starch crops to replace petrol and biodiesel from oil crops. smaller than hydrolysis/fermentation of sugar and starch
The resurgent interest in biofuel at the beginning of the to ethanol). The productivity of some commonly used
twenty-first century is a result of more complex forces. High crops expressed in gasoline equivalents to account for dif-
oil prices caused by increased demand are now combined ference in motive power is presented in the figure below.
with concerns over national energy security and targets to The data reveal that various combinations of physiological
mitigate climate change. Biofuel production requires energy variables can produce high or similar yield and that per-
to sow, grow and harvest crops, including that for machine haps the most important determinant, as seen in both sug-
construction and operation, production and use of agro- arcane and oil palm, is the favourable year-round growing
chemicals, transport of crop products and finally transfor- environment of the tropics. Crops restricted to seasonal
mation into biofuel. Important questions are: First, what are environments produce less, and except for oil palm, oilseed
the energy benefits of biofuel? Second, to what extent can crops are the least productive. Oil palm biodiesel and sug-
biofuel mitigate global warming? Third, how much extra arcane bioethanol alone have high energy efficiencies in
primary production can the biosphere accommodate along the range 8–10 (Liska and Cassman, 2008) while no other
with food requirements for all creatures great and small? crop consistently exceeds 2. Gnansounou and Dauriat
(2005) provided average values for sugar beet (1.83), cereals
Energy efficiency
(1.76) and maize (1.44). While the productivity of soybean
This is the ratio of biofuel output to support energy used
is small, its energy efficiency of 1.9 (Liska and Cassman,
in crop production and subsequent transformation. Plant
2008) is greater than that of maize, presumably reflecting
physiology provides important explanations of compara-
comparative advantages of N fixation and low energy cost
tive productivity and efficiency through climatic adapta-
of trans-esterification to biodiesel.
tion, including C4 photosynthesis to tropical year-round
growing environments (greater production than C3 types
in seasonal environments), metabolic costs of product Greenhouse gas mitigation
type (decreasing in the order oil, starch, sugar), method of A popular view of biofuel use is that since CO2 released
storage (vegetative organs do not depend on successful on combustion was fixed in photosynthesis, impact on
seed set and can accept assimilate over longer periods of the atmosphere is carbon, and hence greenhouse gas,
growth), nitrogen fixation in legumes to reduce the high neutral. That is not the case, however, because fossil fuel
energy cost of nitrogen fertiliser, repeated flowering and is widely used to support production and because crop
fruit set to maintain high sink strength (some crops, e.g. oil production releases oxides of nitrogen and sulphur, and
palm, flower and set seed throughout the year, while most in moist conditions methane, that are more powerful
others just once) and, again related to product type, the greenhouse gases than CO2.

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Biofuel productivity (l ha1 of gasoline equivalent, bioethanol 0.70, biodiesel 0.91) of various crops based on average yields
in countries where they are well adapted. Yields (t ha1) are 8.0 for wheat in the UK, 9.4 for maize in USA, 78 for sugar beet in
France, 74 for sugarcane in Brazil, 2.7 for soybean in Argentina, 2.1 for sunflower in the Czech Republic, 1.7 for rapeseed in
Canada and 21 for oil palm in Malaysia (from FAOSTAT http://faostat.fao.org/default.aspx).
(Continued)
10 CHAPTER 1: Sustainable Agriculture and Crop Physiology

BOX 3  Continued
The relatively few analyses of greenhouse gas balance of The future
biofuel production reveal that it is not always significant. Concern about competition with food production has
While Hill et al. (2006) established a 12% saving over gas- promoted two reactions to sustain the quest for a biofuel
oline for bioethanol from maize, Farrell et al. (2006) could economy: first, utilisation of currently unharvested vegeta-
find none. In contrast, that study reports a gain of 43% tive parts of crops and forests and, second, use of non-food
for biodiesel from soybean. The inevitable conclusion crops, including woody plants. Most target non-food crops
is that even without any greenhouse costs associated will, as do crop residues, provide lignocellulosic vegetative
with land clearing for the expansion of cropping areas material. A biomass crop of current focus is switchgrass
to meet the requirements, associated emissions of CO2 (Panicum virgatum), which has high productivity and wide
from the decomposition of organic matter and microbial adaptability (McLaughlin et al., 1999). A biodiesel favourite
release of nitrous oxides, biofuel does not substantially is Jatropha curcas touted to perform well in marginal areas
benefit greenhouse gas mitigation. Where land is cleared but as yet without evidence. The claims of its productivity
for biofuel crops, greenhouse gas emissions may be dou- on marginal and degraded land (Francis et al., 2005) are
ble those from equivalent gasoline use for 30 years and unlikely to be established. So far, there are no large-scale
remain positive for 167 years (Searchinger et al., 2008). production units from which to evaluate energy efficiency
or economic viability of biomass crops. Experimental data
Competition with food production suggest energy efficiencies of bioethanol from lignocellu-
Marked increases in grain prices in 2007 have revealed losic crops in the range 1.8–5.6 (Gnansounou and Dauriat,
the impact of a relatively small diversion of crop produc- 2005). Further, non-food crops always compete with food
tion to biofuel. These have found support among farm- crops for land, water and nutrients. The hope of using mar-
ers and hope in some development agencies for a much ginal land is fatuous, in a world seeking to increase food
required boost to agriculture in developing countries. production. Land that can grow energy crops could also
Consumers, while expressing concern for the environ- grow food. There is no essential difference.
ment as forestland is converted to biofuel production,
fear the era of cheap food has come to an end. Certainly, Future emphasis on land-based biofuel production is
competition for crop products for fuel and food between likely to concentrate on residues from agriculture and
the rich and the poor has the capacity to further increase forestry, and any level of success will await development
food prices as the demand for food continues to increase. of commercially efficient technology for transformation to
Agriculture now provides, with some shortfalls, food for 6.5 biofuel. It will also require careful site-by-site analyses of
thousand million people and is challenged to do better for how much residue can be removed without deleterious
about 9.0 thousand million by 2050, while conserving the effects on productivity. Residues are not wastes left to rot
resource base and preserving as much land as possible for on the ground but food for domestic animals and a web
nature and its other values. From an agronomic perspec- of consumers and decomposers that play important roles
tive, the doubling of food production during the next 40 in maintaining the natural environment and the physi-
years is accepted by international development organisa- cal and chemical condition of soils. The apparent large
tions as an enormous challenge. Biofuel adds a large extra net energy benefits of these sources will diminish once
burden. The standard nutritional unit of 500 kg grain equiv- they are recognised as part of production systems and
alent per head per year is a convenient way to express appropriate proportions of support energy are allocated
agricultural production in relation to human food supply to them. Biofuel does not, it seems, generate enough
(Loomis and Connor, 1996). The unit is sufficiently large energy to justify the many environmental (Scharlemann
to allow some production to be used as seed for the next and Laurance, 2008; Searchinger et al., 2008) and social
crop, some to be fed to animals, some to be lost in storage, impacts of large-scale production. The most likely out-
and some land to be diverted to fruit and vegetable crops. come is that hopes for a land-based biofuel economy will
Significantly, that same amount of production is sufficient fade. Evidence suggests a modest (10%) contribution to
to produce just 200 l of maize ethanol (2.5 kg l1), enough national motor fuel supply for countries with large crop-
for 2000 km at 10 l per 100 km (Connor and Minguez, 2006). land resources relative to population size but that biofuel
The carrying capacity of land for cars is just too small and cannot be a major part of a solution in a highly populated
increasing population must take precedence for food sup- and energy-demanding world. The energy now used
ply. Even dedicating all US corn and soybean production to generate biofuel, especially natural gas in ethanol
to biofuel would provide only 12 and 6%, respectively, of production, may well find more efficient use directly in
the current national demand for gasoline and diesel. transport.
 5. Aims and Structure of This Book 11

5. Aims and structure of this book

Agriculture operates in the context of production and profit, environmental and social outcomes and food
quality and safety (Section 2). The world faces challenges related to the increasing demand of food and
other agricultural products (Section 3), competing land uses, and misplaced emphasis on ecologically, eco-
nomically or socially unsound agricultural practices (Section 4). Against the twin framework of the current
paradigm of agriculture, and the interactions between biophysical sciences contributing to agricultural pro-
duction, the primary aim of this book is to highlight the value of crop physiology as an interface discipline
whose practical benefits are realised by targeted links with genetic improvement and agronomy. Whenever
suitable, this book seeks to draw attention to and partially counteract the reductionist views more common
in disciplines targeting lower levels of organisation; we will emphasise conserved traits, where definitions at
low levels of organisation remain relevant at the crop level (e.g. phenological development, disease toler-
ance and herbicide resistance), as distinct to those traits that are typical of populations, for which relation-
ships with lower levels of organisation are not straightforward (e.g. grain yield).

This book comprises three parts: Part 1 focuses on selected, contrasting farming systems of the world to
sample more specific challenges of agriculture, Part 2 outlines principles of capture and efficiency in the use
of crop resources, and Part 3 concentrates in recent and projected advances in breeding and agronomy with
emphasis on physiological perspectives, scale issues or both.

5.1. Part 1. Farming systems: Case studies

Whereas, globally, the challenges of sustainable agriculture are similar, the particular drivers and constraints
of specific farming systems can be vastly different. Part 1 thus contrasts predominantly rainfed systems –
including the cereal-based systems of Australia (Chapter 2), where rainfall patterns constraint and shape
farming options; the diverse farming systems of the Pampas of Argentina (Chapter 3), where dramatic inten-
sification of agriculture seeks to improve the capture of resources, chiefly rainfall, dilute fixed costs and
maintain the resource base; and short-season, cereal-based systems of Finland, where low temperature
and long days are major drivers – whereas EU regulations set strict guidelines for agriculture (Chapter 4).
Part 1 closes with an overview of diverse, cereal-based farming systems of Asia, ranging from less than
1 t ha1 year1 for sole crops in dry lands to 15 t ha1 year1 in irrigated intensive systems, allowing for mul-
tiple cropping (Chapter 5).

5.2. Part 2. Capture and efficiency in the use of resources:

Quantitative frameworks
The shift from plant growth analysis based on relative growth rate and its components, that is, net assimila-
tion rate and leaf area ratio, to crop growth analysis based on capture and efficiency in the use of resources
was a milestone in crop physiology (Monteith, 1994). Part 2 of this book focuses on capture and efficiency
in the use of water, radiation and nitrogen with a strong quantitative perspective. Chapter 6 has a dual aim:
It considers the challenges of irrigated cropping systems worldwide, hence expanding the focus on rainfed
agriculture developed in Part 1, and accounts for water productivity from engineering to crop scales. Chapter
7 outlines the use of radiation-based methods to analyse crop growth in agronomy, breeding and model-
ling, and Chapter 8 presents a comprehensive physiological perspective of nitrogen nutrition in field crops
and pastures.

For updates on other important nutrients not considered in this book, chiefly phosphorus, readers are
referred to Lambers et al. (2008), van der Heijden et al. (2008) and Sawers et al. (2008) for recent accounts
of nutrient acquisition from evolutionary and ecological perspectives that emphasise the role of mycorrhiza
and other soil organisms; Bucher (2007) for a closer look at the biological aspects of phosphorus uptake
12 CHAPTER 1: Sustainable Agriculture and Crop Physiology

at the root and mycorrhiza interface; and reviews with an agricultural focus by Ulen et al. (2007) and
Hart et al. (2004). Whereas nutrients have generally been considered individually, for example, manage-
ment of nitrogen or phosphorus, integrated approaches accounting for interactions between nutrients are
attracting more attention (Dobermann et al., 1998; Kho, 2000; Sadras, 2006; Saïdou et al., 2003; Yadav
et al., 2000).

5.3. Part 3. Crop physiology, breeding and agronomy

The focus of this book is the crop, which biologically is a population with both properties that depend on
its constitutive individuals, and emergent properties that are unique to this level of organisation (Donald,
1981). The insightful perspective of Güiraldes opening this chapter highlights the dual feature of the popu-
lation: It depends on its constitutive individuals, but it is also an entity on its own, with properties that are
unique to this level of organisation. Furthermore, relevant attributes of farming systems involve community-
level processes where the crop species are primary producers, alongside with weeds, closely interacting with
organisms up the trophic chain, including pathogens and decomposers.
A mature crop physiology discipline has grown out of its own, early attempts to increase productivity
through a naive, linear scaling up across levels of organisation, but this syndrome is still common in other
disciplines (Hammer et al., 2004; Sinclair et al., 2004). Lack of appreciation of both trade-offs and buffer-
ing processes involved in scaling up and crop-level attributes that override seemingly sound traits at lower
levels of organisation remains an issue and a major source of inefficiency in the investment of research effort
(e.g. Vinocur and Altman, 2005; Yamasaki et al., 2005; Zhu et al., 2007). Figure 4 contrasts a general framework
of biotechnology and crop physiology. Where biotechnology focuses on very detailed molecular mecha-
nisms at the expense of environmental factors and higher organisation levels, crop physiology focuses on
the population level and attempts to integrate across levels of organisation, with greater emphasis on the
nuances of environmental drivers of crop growth, development and yield formation.
Part 3 deals with the links between breeding, agronomy and crop physiology against the conceptual frame-
work outlined in Figure 4 (right). Chapter 9 presents an evolutionary perspective that, we propose, provides
the theoretical ‘glue’ for our conceptual model, whereas Chapter 10 updates efforts aiming at its mathe-
matical formalisation. Biological phenomena can be explained by invoking proximal, physiological mecha-
nisms and ultimate evolutionary causes. Both approaches are valuable, and they are not mutually exclusive.
The photoperiodic response of flowering plants can be explained physiologically, from gene to phenotype
(Chapter 12). However valuable, this explanation is incomplete, as the basic question of why wheat is a
long-day plant remains unanswered. The answer to the ‘why’ question requires evolutionary considerations,
which are often speculative but nonetheless illuminating. There is a rich history in crop science demonstrat-
ing the insight gained from the consideration of evolutionary aspects of individual plant traits (Denison
et al., 2003; Donald, 1968, 1981; Donald and Hamblin, 1983; Evans, 1984, 1993; Troyer, 2006). Chapter 9
explores the notion that trade-off-free options to increase competitiveness (e.g. increased gene expres-
sion) have been comprehensively tested by natural selection. This leads to the concept that further genetic
improvement of crop yield would therefore involve substantial trade-offs between fitness in past versus pres-
ent environments, between individual competitiveness and the collective performance of plant communities
or between individual and multiple crops, or a combination of these trade-offs.
There is an inherent limitation in adaptive explanations dissecting organisms into unitary traits (Gould and
Lewontin, 1979), but recent developments in modelling crops from gene to phenotype allow for integration
of individual traits and to capture emergent properties at the individual and population levels (Chapman,
2008; Hammer et al., 2006; Hoogenboom et al., 1997, 2004; Messina et al., 2006). Chapter 10 presents
the state-of-the art in this new generation of mathematical models used to explore the links between crop
physiology and the critical interaction between genotype and environment, with a proclaimed potential
to help integrating molecular technologies in plant improvement. Chapter 11 expands on physiological
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