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 Future Implications for Animal Production: A Perspective on Sustainable Livestock
Intensification1
Luis Orlindo Tedeschi1, James Pierre Muir2, David Greg Riley1, Danny Gene Fox3

1 Department of Animal Science, Texas A&M University, College Station, TX, USA
2 Texas A&M AgriLife Research, Stephenville, TX, USA
3 Department of Animal Science, Cornell University, Ithaca, NY, USA

Abstract. Food supply has improved considerably since the 18th Century industrial revolution, but
inadequate attention has been given to protecting the environment in the process. Feeding a
growing world population (estimated by the United Nations to be 9.55 billion by 2050) while
maintaining or reducing the impact on the environment requires immediate and effective solutions.
Sustainability is difficult to define because it embodies multifaceted concepts and the combination
of variables that make a production system sustainable can be unique to each production situation.
The concepts of sustainability, resilience, and resistance are dependent upon context and time
horizon. Sustainable is the characteristic of a system that continues to co-exist with other systems
at a different output level after a period of perturbation(s). Resilience is the ability of a system to
fully (or partially) recover and re-establish a steady output (i.e., dynamic equilibrium) after it has
been perturbed either by a natural (i.e., endogenous) or an artificial (i.e., exogenous) stressor.
Sustainable intensification (SI) produces greater output(s) through the more efficient use of
resources within a period of time while reducing (or at least not increasing) negative impact on the
environment; hence, SI provides opportunities for increasing animal and crop production per unit
of area via sustainable alternatives that fully consider the three pillars of sustainability: planet,
people, and profit. The question that remains unanswered is whether global agriculture in 35 years
will feed 10 billion of people, produce biofuel and clean electricity, supply fiber, and at the same
time reduce its environmental footprint to avert or even reverse climate change.
Key words. Cattle, Efficiency, Environment, Production, Sustainability, Systems
The Need for More Productive and Sustainable Food Production Systems
Discussions about sustainability come at a time when the awareness of global warming and
food shortage risks has never been greater; these discussions were brought about by recent extreme
weather events and hunger around the world. Frustration with the lack of achievements in the
prevention of global warming was augmented by the perception that the 2009 international climate
negotiations in Copenhagen failed miserably; it was a “festival of conspiracy and betrayals”
(Goodell, 2010). The sustainability issue could go in the same direction and become entangled in
the same web of politics as the Copenhagen negotiations if science is not taken seriously by
conquering environmental pollution and hunger simultaneously. China blames the United States
of America and Europe for global warming, saying that it has been largely caused by 200 years of
fossil-fuel burning, after the industrial revolution in the 1700s (Goodell, 2010). Clearly, this is not
a one-player game; it requires multilateral decisions and measurement on a global scale.
In an unprecedented move by the Catholic Church, the concern of global warming
threatening life on Earth was the central point of the encyclical “Laudato Sí” (i.e., praise be to you)
by the Pope Francis: “a very solid scientific consensus indicates that we are presently witnessing

1
Paper presented at the 52th Annual Meeting of the Brazilian Society of Animal Science, Belo Horizonte, MG on July
19-23, 2015 (http://www.sbz2015.com.br/). Citation: Tedeschi, L. O., J. P. Muir, D. G. Riley, and D. G. Fox. 2015.
Future implications for animal production: A perspective on sustainable livestock intensification. Pages 1-23 (CD
format) in Proceedings of the 52th Annual Meeting of the Brazilian Society of Animal Science, Belo Horizonte, Minas
Gerais, Brazil. Sociedade Brasileira de Zootecnia (SBZ)

1
a disturbing warming of the climatic system” and emphasized that “nobody is suggesting a return
to the Stone Age, but we do need to slow down and look at reality in a different way, to appropriate
the positive and sustainable progress which has been made, but also to recover the values and the
great goals swept away by our unrestrained delusions of grandeur” (Francis, 2015). Global
warming has become a political and religious matter, but what does science tell us?
Environmental protection is paramount for providing minimum livelihood standards for
humans and ensuring the survival of our species (and many others) in centuries to come. This
spans the preservation of biodiversity and responsible stewardship of soil and water resources as
we seek to feed and clothe ourselves utilizing these natural resources. Widespread alterations of
the environment by humankind in support of our own existence have accelerated since the
beginning of the industrial revolution in the 18th Century (Sabine et al., 2004). Food production
has been based on maximizing productivity and profitability with inadequate concern for
protection of soil, water, and water quality in the process. As a result, these alterations have
disrupted natural cycles by adding sequestrated carbon back into the atmosphere at a much faster
pace than it can be immobilized.
Terrestrial ecosystems react to atmospheric CO2 concentration such that their short- and
long-term feedbacks likely play a role in climate change. Their precise contribution is largely
uncertain because they impact diverse ecozones and the time of impact cannot be predicted.
Although uncertainty remains, Schimel et al. (2015) reported a significant uptake by tropical
forests and suggested that up to 60% of their present-day terrestrial sink is caused by increasing
atmospheric CO2. On a global scale, the FAO (2006) reported that the livestock sector was
responsible for 18% of total greenhouse gas (GHG), when expressed as CO2 equivalent, and 9%
of anthropogenic CO2 emissions. These estimates, however, have been challenged and thought by
some to be significantly less (Hristov et al., 2013; Place, 2015).
More than 215 years ago, Thomas Robert Malthus prophesied that human population
growth would outrun our ability to produce food (Wrigley, 1988). Today this remains a real
possibility given the need to feed a staggering growth in world population, projected to be 9.55
billion by 2050 (United Nations, 2013), while maintaining or reducing the impact on the
environment. This task poses a progressively more challenging and constant pressure on crop, soil,
and animal scientists around the world to come up with immediate and effective solutions. Our
modern food production involves intricate and complex systems with many feedback signals
(Tedeschi et al., 2011b). For instance, an increase in food production affects the environment by
changing the climate directly through emissions that contribute to global warming and from excess
nutrients that reduce water quality or by reducing nonrenewable resources (e.g., fossil fuel). In
turn, the changes in environment offset improved productivity through harsher conditions such as
degradation of soil quality, increase in warming, resurgence of new diseases, depletion of
biodiversity, and loss of adapted animals and crops, among many other outcomes. Animal
scientists must develop strategies that forecast the rate and magnitude of global changes as well as
their possible influences on the food production chain. They cannot, however, afford to stop there:
they must then develop tactics to adapt to and mitigate the causes of global climate change due to
food animal production.
The Feed the Future2 concept likely had its inception in the 1960s, but not until recently
has it been given much attention. The success of the Feed the Future initiative depends on
innovative solutions to foster greater agriculture production while maintaining profitability and
minimizing its environmental impact. The current stagnant growth rate of food animal agriculture
will likely fail to meet the expected growth in global demand for animal protein, especially if it is

2
http://www.feedthefuture.gov

2
to be profitable when produced in an environmentally and socially responsible way. Innovations
and investments are needed in animal science research and development in the 21st century as
dramatic increases in global demand for food protein (e.g., meat, fish, eggs, and dairy products)
are forecasted by 2050 due to increase in world population (National Research Council; NRC,
2015). Additionally, the possible use of biomass for biofuel production through thermochemical
conversion technologies (Verma et al., 2012) creates a new, imminent threat to livestock systems
because they often compete for the same land area and resources. The future of this competition is
uncertain as it depends on regulations, subsidies, and petroleum prices, and these policies vary
across countries (Hertel, 2011).
Debates on global warming and Feed the Future concepts have initiated unprecedented
discussions about alternative production systems needed to meet growing human demand for food.
Unprecedented extreme high temperatures (> 40oC) and the number of days with temperature
humidity index above comfort threshold (i.e., 68) have noticeably increased in recent years in
countries located within the temperate zone, thereby negatively affecting agriculture and livestock
productions (Silanikove and Koluman, 2015). Regardless of the disagreements about the veracity
and acceptability of global warming data and the methods used to collect it, the weight of the
evidence favors acceptance rather than rejection and we cannot afford to take the risk of being
wrong. Thus, to avoid mass starvation in the future, we must assume there will be continued
exponential growth of the human population (United Nations, 2013) and current methods to
produce food must improve to meet this challenge, using environmentally-friendly technology.
Although many of these technologies exist, they may only partially address the Feed the Future
promise without reducing environmental impact, even if fully implemented. For instance, animal
agriculture expansion into new lands would likely not be a feasible solution because of resulting
increased competition for land from other human activities (e.g., row cropping, urbanization or
water capture; Pretty et al., 2011) among many other issues. The NRC (2012) indicated that “food
security for all” must be achieved in a sustainable way to be achieved long term.
Sustainability Concepts and Definitions
The increase in concerns about sustainable food production systems has resulted in
rhetorical discussions and diverse opinions on what sustainability is. Opinionated papers have
presented contrasting schools of thought (e.g., business-as-usual optimists, environmental
pessimists, new modernists to name a few; Pretty, 1997). Open discussions based on sound science
that lead to clearly defining and describing sustainable food production systems under different
plant and animal production conditions are needed. As humans have imposed greater demands
upon natural systems (e.g., agriculture), some have raised alarming concerns about the
sustainability of these systems because of the finite nature of soil and water resources (Arrow et
al., 1995; Meadows and Meadows, 2007; Meadows et al., 1972) and the heavy dependency of
agriculture on non-renewable resources such as fossil fuel (NRC, 2010). In fact, the 2007-2008
commodity crisis highlighted food production vulnerability to weather-related events, financial
markets, and poor governmental interventions in protecting food exportation to avert chaos in the
domestic food supply (Hertel, 2011).
A recent concept, Sustainable intensification (SI), accounts for the need to increase crop
and food animal production per unit of area while taking into consideration sustainable production
alternatives that fully address the three pillars of sustainability (planet, people, and profit; Makkar
(2013)). Thus, SI requires that food production systems be profitable, socio-culturally acceptable,
beneficial to the people, as well as environmentally friendly to natural resources (Figure 1).
Sustainable intensification supports climate change mitigation and the Feed the Future initiative.
Improving food yields in areas that are already highly productive is not necessarily the
solution because it could end up increasing the environmental burden. For about 80% of the

3
chronically hungry in Africa, most of whom are smallholder farmers, an improvement in yield per
area would increase their access to food and generate more income (The Montpellier Panel, 2013).
In that context, SI is used to increase productivity in areas that were previously underutilized. A
2010 report analyzing 40 SI projects from 20 countries during the 1990s to 2000s indicated that
about 10 million farmers had benefitted from SI projects with improvements on approximately
12.75 million Hectares (Pretty et al., 2011). Animal products and crop yields per hectare were
increased by combining the use of new and improved crop varieties. The main challenge with the
adoption of SI is the spread of effective processes and lessons to many more people, generally
smallholder farmers and pastoralists across the world (Pretty et al., 2011).

Figure 1. The three pillars of sustainability. Based on “sustainable development” from

Wikimedia.org under Creative Commons licensing, and further adapted from United Nations
(1987), International Union for Conservation of Nature (IUCN) (2005), Makkar (2013), and
Makkar and Ankers (2014).

Unfortunately, the deployment of SI technologies does not occur without controversy,

which is often based on incomplete information or lack of sound science. An example of SI
technology conflicting with social issues is the misperception of feeding genetically modified
(GM) plants or their byproducts to animals. It is generally accepted by the scientific community
that GM plants (e.g., corn) used as feed material for food-producing farm animals are safe because
they do not alter animal metabolism or the organoleptic and bromatological characteristics of meat,
milk, and eggs (Swiatkiewicz et al., 2014). The negative perception of GM food by some, however,
hinders its scope and application where it is needed the most, and limits its contribution to SI. The
GM organism can be a powerful tool in addressing climate change issues as discussed by
Zilberman (2015): the first generation GM plants are insect and disease resistant or herbicide
tolerant; the second generation GM plants have enhanced organoleptic and bromatological
characteristics, photosynthetic efficiency, and abiotic stress tolerance (e.g., drought); and the third
generation GM plants produce specific chemicals and pharmaceuticals.

4
 Another social conflict with SI stemming from misconception is that some believe
livestock always competes with human food supplies because grain (e.g., corn) or cultivated
pasture used to feed livestock could be used directly as human food and, thus, they assume
livestock activity is an inefficient or wasteful use of resources (CAST, 2013). In actual fact, as
shown in Table 1, for some species such as beef and dairy cattle, the protein conversion efficiency
on a human-edible basis is often greater than 1:1, indicating that the amount of human-edible
animal protein product is greater than the human-edible feed consumed by the animal (CAST,
1999), confirming the undeniable benefit of animals as an efficient source of high-quality food for
humans. Thus, the dilemma in hand is how to meet the challenge of providing high-quality food
to a growing human population while reducing the environmental footprint in an effective,
economical, and timely fashion.

Table 1. Comparative gross efficiencies of conversion of dietary energy and protein to product
and returns on human-edible inputs in products for swine, poultry, milk, and beef in different
countries 1
Country Species Gross efficiency Human-edible efficiency
Energy Protein Energy Protein
Argentina Swine 0.15 0.07 0.24 0.11
Poultry 0.18 0.30 0.28 0.69
Milk 0.19 0.16 4.61 1.64
Beef 0.02 0.02 3.19 6.12
Mexico Swine 0.13 0.08 0.25 0.21
Poultry 0.2 0.33 0.34 0.83
Milk 0.12 0.11 0.79 1.06
Beef 0.06 0.02 16.36 4.39
South Korea Swine 0.20 0.16 0.35 0.51
Poultry 0.21 0.34 0.30 1.04
Milk 0.26 0.19 3.74 14.3
Beef 0.06 0.06 3.34 6.57
United States Swine 0.21 0.19 0.31 0.29
Poultry 0.19 0.31 0.28 0.62
Milk 0.25 0.21 1.07 2.08
Beef 0.07 0.08 0.65 1.19
1 Gross efficiency was calculated as outputs of human-edible energy and protein divided by total

energy and protein consumed by the animals. Human-edible efficiency was computed as outputs
of human-edible energy and protein divided by human-edible energy and protein consumed by the
animals.
Adapted from Council for Agricultural Science and Technology (1999).

Since the introduction of the sustainable development concept, which was defined as
“development that meets the needs of the present without compromising the ability of future
generations to meet their own needs” by the World Commission on Environment and
Development (United Nations, 1987 -- Brundtland Commission's Report), sustainability has been
defined in more than 100 different ways (Pretty, 1997) probably because one definition does not
fit all possible scenarios. Sustainability fits the criteria for a “wicked problem” (Peterson, 2011):
(1.) the ideal definition lacks specificity and it can be reduced to a slogan, (2.) one can never know
if sustainability has been achieved because the outcome is usually better or worse rather than true

5
or false, (3.) stakeholders have different points of references for the problem, and (4.) system
components and cause/effect relationships are uncertain or changing. Thus, there is not one best
definition of sustainability. Pretty (1997) believes that sustainable agriculture is more a learning
process than prescribed specific and rigorous application of technologies, practices, or policies.
Regardless of the definition(s) adopted for sustainability, one needs to clarify what is being
sustained, boundaries of the system, intensity of disturbances, and time scale. Figure 2 graphically
illustrates some concepts to establish the basics for effective communication and understanding
among parties. It shows different response behaviors when a system (or organism) is challenged
with a temporary (or permanent) perturbation or distress with an onset at time 10. Except for the
unviable scenario portrayed in Figure 2, the other scenarios in general represent different degrees
of sustainable behaviors from an agro-ecological sciences perspective, as defined below.

120
Relative Output Level or Resource Outflow, %

100

80

60 Perturbation

40

Unviable Resilient
20
Sustainable Stable
Responsive
0
0 5 10 15 20 25 30 35 40 45 50
Relative Time

Figure 2. Schematic illustration of different levels of hypothetical sustainable responses by a

system after a period of perturbation(s) or stress(es).

Definitions for sustainability abound in the agro-ecological sciences. The Oxford English
Dictionary defines sustainable as "capable of being maintained or continued at a certain rate or
level," but it does not make any attempt to include in this context the system reaction to a
perturbation. A broader concept of sustainability needs to take into account the context (e.g.,
variables involved and their relationships) and the time horizon. Sustainable is the characteristic
of a system (or an organism) to continue to co-exist with other systems (or organisms) at a different
output level after a period of perturbations. Furthermore, a sustainable system has to maintain itself
with minimal or no impact outside its boundaries. Therefore, one has to establish the chronological
and/or spatial boundaries of the system. Sustainability is the ability of a system to be sustainable.
As illustrated in Figure 2, after a period of perturbations (or distress), the system (or organism)
reduces its output to a lower level and stabilizes at this new level. Sustainability could be difficult
to achieve when exogenous stresses are constantly imposed onto a system, possibly leading to an
eventual collapse.

6
 Resilience is defined by the Oxford English Dictionary as "the quality or fact of being able
to recover quickly or easily from, or resist being affected by, a misfortune, shock, illness, etc.;
robustness; adaptability" or "the action or an act of rebounding or springing back; rebound".
Resilience, thus, is the ability of a system (or organism) to fully (or partially) recover and re-
establish a steady output (i.e., dynamic equilibrium) after it has been perturbed either by a natural
(i.e., endogenous to the system) or an artificial (i.e., exogenous to the system) stressor(s). In other
words, resilience is achieved through mutual reinforcing feedback loops that permit the system to
persist overtime despite temporary disruptions, by either intrinsic (e.g., endogenous) or
surrounding environmental factors (e.g., exogenous) to the system, by regenerating itself to its
original state. Resilience can have a positive or negative impact on the system (or organism)
output. A system is resilient when it recovers to the same level at which it functioned prior to the
perturbation. Under initially unfavorable grazing conditions, a growing animal’s subsequent
compensatory growth is an example of resilience. Animals that have undergone an undernourished
period usually lose body weight (BW) (in the form of body reserves) in support of important bodily
functions to ensure survivability, and then, after the nutritional restriction has ceased, they partially
or fully restore their BW to the normal growth curve by compensating the BW loss via increased
growth rate (Ryan, 1990). Resiliency is the ability of a system (or organism) to be resilient or to
absorb stress without suffering modifications. As illustrated in Figure 2, after a period of
perturbations (or distress), the system (or organism) reduces its output, but immediately initiates
the recovery to its original state.
A sustainable system is resilient within a given context but not all resilient systems are
sustainable because in the process of rebounding, the system may overshoot and collapse, find
another point of temporary stability, or collapse entirely. Therefore, sustainability is not achieved
if the system cannot return to its original stable level. The point of no return occurs when a system
(or organism) can no longer be resilient nor sustainable, and the fate of the system (or organism)
is a general failure as illustrated by the unviable scenario in Figure 2. For example, the mechanism
of compensatory growth has been well documented in grazing animals. It depends on the degree
and duration of the nutritional restriction and the quality of feed provided after the cessation of the
undernourishment period (Ryan, 1990). Some animals can partially or fully restore their
embedded, genetic potential for growth while others may stay at a lower growth or yield level (i.e.,
stunted animals) and never reach their normal growth (Hogg, 1991) even if previous ADG is
reestablished.
Stable is defined by the Oxford English Dictionary as “not likely to change or fail; firmly
established” while resistant is “offering resistance to something or someone.” A stable (or
resistant) system (or organism) is able to produce and reproduce under periodic or continuous
stress conditions for a given time compatible with its growth cycle (e.g., fertilization, development,
yield, offspring, or seeds). The system (or organism) is insensible to perturbations in the
surrounding environment and it maintains an output in dynamic equilibrium over time and across
several generations. In short, it has the ability to resist disorder. As with the previous discussion,
we assumed that the perturbation or distress is within reasonable physiological limits, most notably
non-fatal. As illustrated in Figure 2, after a period of perturbations (or distress), the system (or
organism) does not change its output level. In biological sciences, this outcome is rarely observed,
as most living organisms will respond to an exogenous stressor in either a negative or positive
way. Tolerance or survivability, i.e., the ability of an organism to stay alive during stress conditions
with minimum or absent growth and proliferation, waiting for the right time to express full genetic
potential, is more related to resilience than resistance. For instance, Volaire et al. (2014) defined
drought survival as the ability of plants to cease growth during moisture shortages but to regrow
(continue their life cycle) when drought ceases. Plants, therefore, decrease output level for a period
of time and then recover growth rates (but less likely total yield vis-à-vis their genetic potential)
to pre-stress levels.

7
 Others have defined natural systems as resilient when they tend to maintain their integrity
under disturbances and defined stability as the ability of a system to return to an equilibrium state
after a temporary disturbance (Holling, 1973). These definitions are somewhat opposite to our
definitions as discussed above and shown in Figure 2. Ludwig et al. (1997) used Holling’s (1973)
definitions of resilience and stability and compared them from a mathematical perspective. In the
long run, resilient organisms or systems tend to resist exogenous perturbations by always reverting
to the original level, thereby behaving more like a stable/resistant system (Figure 2). Our
definitions of resilient, sustainable, and resistant are more in line with those adopted by Fiksel
(2003). Our observations support the view that true stability is nearly impossible in natural
ecosystems (or organisms) because disturbance (stress) is almost always an integral part of those.
An unviable system (or organism) is neither sustainable nor resilient to perturbations and
complete failure is the result after a perturbation unless an exogenous interference rescues the
system (or organism) in time. Row cropping annual grains with tilling, irrigation, and other heavy
inputs is an example of such system dependence on exogenous interference. Even at that time,
however, the system (or organism) may or may not find a new level of stability.
We conclude SI is the best overall concept that describes the approach that will be needed
to increase food production while accounting for environmental effects. SI is about producing
more output(s) through the more efficient use of the resources (i.e., inputs) within a period of time
while reducing the negative interference with the environment. The origin of SI dates back to
1990s when the term was developed for African production systems in which low yields and high
environmental degradation were predominant, prompting an immediate pejorative use of SI to be
synonymous with smallholder-oriented or organic production technology (Garnett and Godfray,
2012). SI has often been confused with other terms and concepts such as conventional
intensification, ecological intensification, agroecology, organic agriculture, climate smart
agriculture, and eco-efficiency (Garnett and Godfray, 2012; Kuyper and Struik, 2014) that have
triggered negative perceptions. Tittonell (2014) concluded that SI is more loosely defined than
ecological intensification. Struik et al. (2014) indicated that SI is by definition an oxymoron
problem as a win-win situation is rare, and ambiguities exist in both terms of SI (Uphoff, 2014).
We agree with The Royal Society (2009) definition of SI, disconnected from any particular
agricultural production system, as a form of production wherein “yields are increased without
adverse environmental impact and without the cultivation of more land.”
Campbell et al. (2014) concluded that SI and climate-smart agriculture are complementary
processes but different multi-managerial approaches have to be adopted at different levels to
achieve successful implementation. These include diversified farming systems, local adaptation
planning, building responsive governance systems, enhancing leadership skills, building asset
diversity, reducing consumption and waste, building social safety nets, facilitating trade, and
enhancing diets. It is, however, important to stress that SI entails the increase of food production
from existing farmland without additional environmental impact. The responsive behavior shown
in Figure 2 illustrates SI: output level increases after the onset of perturbations in the system until
it reaches a dynamic equilibrium at a greater level than before the perturbations. Of course, in this
case, perturbations had a positive impact on the system (or organism), assuming that the level of
input(s) were constant.
Sustainable production is often misunderstood as a goal for farmers and other land
managers. Sustainability represents the state of a complex dynamic system that is always evolving
and transforming itself; thus, it cannot entail a production goal. It is an intrinsic characteristic of
the system that needs to be maintained. The system has to be shaped and managed until it is
sustainable. For instance, sustainable profitability and SI are distinct means to be productive in
distinctive ways: the former contributes to sustainability by decreasing cost-to-benefit ratios while
the latter one does so by increasing the output per unit of input. A systems approach incorporates

8
multiple means to attain system sustainability. For example, input resource use and energy
intensity reduction, waste conversion into valuable products, identification of boundaries,
establishing requirements, and employing feasible technologies are among key strategies for
system sustainability (Fiksel, 2003). In addition, from a systems perspective, we must identify key
processes within the system of interest and define their dimensions (Uphoff, 2014). Furthermore,
while SI may improve food supply in the foreseeable future, it does not imply food security for all
(Garnett et al., 2013) because food availability and supply, distribution and allocation, storage, and
utilization may prevent food from reaching those that need it the most (NRC, 2012). In fact,
nutrient security might be a better term for addressing hunger.
Fath (2014) believed that we can hardly address sustainability of ecological systems
without discussing ideas about systems thinking put forward by three personalities: the American
ecologist Bernard Patten, German sociologist Niklas Luhmann, and Austrian-born architect
Christopher Alexander. Patten’s (1978) contributions to the importance of defining environment
in ecological systems and Luhmann’s (1996) for social systems indicate that boundaries are
necessary to demarcate the limits of the system and the environment. Boundaries are also key to
understanding how systems interact with environment in exchanging energy, matter, and
information that keeps the system active and sustainable. The parallel insights of these individuals
remarkably emphasizes the ideas discussed by Forrester (1961, 1971, 1973), Sterman (2000),
Maani and Cavana (2007), Morecroft , and Warren (2008) for business dynamics, and by Tedeschi
et al. (2011b) for animal agriculture on using a systems approach when modeling complex
relationships, including education (Lander, 2015).
The concepts and definitions discussed above lay the foundation for studying SI systems
and proposed ways to describe, measure, and evaluate their achievements. Methods to effectively
promote SI as well as the deployment and stability of SI systems need to be addressed. The NRC’s
(2010) “Toward Sustainable Agricultural Systems in the 21st Century” assessed the scientific
evidence of strengths and weaknesses inherent in deploying sustainable agriculture under various
production, marketing, and policy approaches. It also examined its unintended consequences. The
NRC (2010) committee identified incremental and transformative changes. The latter are critical
and include “the development of new farming systems that represent a dramatic departure from
the dominant systems of present-day American agriculture and capitalize on synergies and
efficiencies associated with complex natural systems and broader social and economic forces
using integrative approaches to research and extension at both the farm and landscape levels.”
An important practice for livestock suggested by the committee was the genetic improvement of
livestock to increase feed efficiency as well as animal health and welfare.
Sustainable Livestock Intensification (SLI)
Livestock production impacts the environment in many ways. Janzen (2011) indicated that
about 20 to 30% of land utilized by humans is used for grazing while another 7 to 10% is dedicated
to producing animal feed and forage. A large portion of grazed areas is not suitable for alternative
production of human food. In the United States, about 25.9% of the land is classified as grassland,
pasture, or rangeland (CAST, 2012). Oltjen and Beckett (1996) indicated that humanly edible
returns for digestible energy ranged from 37 to 59% and returns to digestible protein ranged from
52 to 104%, depending on the time spent in the feedlot and the amount of corn fed. Similarly,
Wilkinson (2011) indicated that feed conversion ratio for edible protein into edible animal protein
was greater than 1. Although certain ruminant production systems have a human-edible protein
efficiency greater than 1 (Table 1) (CAST, 1999), about 16,000 L of water is required for each
kilogram of beef produced, though there is large variation in that estimate (Janzen, 2011). The
main source of agricultural N loss to air and considerable N and P emissions to aquifers and surface
water originates from livestock (Janzen, 2011); thus, sustainably managing livestock N and P

9
excretion (Eghball, 2002; Klausner et al., 1998), especially in concentrated animal feeding
operations, is necessary to contain excessive environmental pollution.
Livestock provide several benefits, including the conversion of human-inedible feed into
high-quality food (Gill, 2013), preservation of ecosystems (i.e., grasslands), recycling of organic
matter and nutrients, and many social aspects associated with livestock (Cheeke, 1999; Janzen,
2011). We therefore need to make livestock production more sustainable using novel technology
and rational management strategies while simultaneously providing high-quality food, preserving
biodiversity, and adopting animal welfare practices. Sustainable livestock intensification is needed
in animal production through the use of technologies that can be used to increase food from
animals. Dumont et al. (2013) proposed five ecological principles for the redesign of animal
production systems (management to improve animal health, decreased inputs, optimize metabolic
functioning of farming systems, enhance diversity, and preserve biological diversity) and Dumont
et al. (2014) discussed 40 issues that were collapsed into four major themes: animal adaptive
capacities, feed resources and forage systems, design and evaluation of new production system,
and rules for scaling-up agroecological animal production systems. We believe that focus areas
include management of grassland/row crop fields, precision feeding, adaptation of livestock to
climate changes, water usage and recycling, and the preservation of biodiversity through genetic
manipulation and preservation.
Smallholder animal farmers and SLI
Strategies for intensifying ruminant production around the world cannot ignore smallholder
or subsistence farms and herders. For decades, livestock in the tropics have held the greatest
promise but also the strongest challenges (Preston, 1990), a situation that has only intensified with
the realization that the vast majority of these smallholders exist in developing countries (Kruska
et al., 2003) and are most vulnerable to climate change (Musemwa et al., 2012). Animal health
(Suriyasathaporn, 2011; Young et al., 2014), reproduction (Bahmani et al., 2011), nutrition
(Atuhaire et al., 2014; Stür et al., 2002), household food security (Nampanya et al., 2014), and
profitability (Widiati et al., 2012), especially as it relates to market access (Zvinorova et al., 2013),
are hurdles to overcome if SI is to reach the majority of the world’s ruminant producers. Ineffective
extension, inadequate education, inaccessible credit and gender inequality for women participants
keep progress to a minimum (Esilaba et al., 2005; Fon Tebug et al., 2012; Mekonnen et al., 2010;
Zvinorova et al., 2013). The picture is not all negative, however. Understanding the whole farming
system from a human as well as a biological perspective can make a difference, as does the
involvement of smallholders in identifying bottlenecks and solutions (Bayemi et al., 2009; Esilaba
et al., 2005; Le Gal et al., 2013; Mekonnen et al., 2010; Stür et al., 2002).
The role of decision support systems to assist SLI
The lack of awareness and limited knowledge of mathematical models are the main factors
that foster a negative perception of modeling and simulation, thereby hindering their development
and broader application. Mathematical models have a crucial role in shedding light on unforeseen
variable relationships and quantifying expected outcomes resulting from alternative decisions in
the production scenarios given the context for which the model was intended to be used. Our
reasoning is endorsed by the recommendations of Garnett and Godfray (2012), who recommended
that a more system-oriented approach to decision making is needed to develop substantial
programs of future activity related to SI.
Recently, the outbreak of Ebola in West Africa required real-time modeling and simulation
to identify the spread of the disease and to provide timely guidance for policymakers (Lofgren et
al., 2014). Mathematical models might also be an effective tool to circumvent our imperfect ability
to detect disease outbreak in livestock (Perry et al., 2013) within the SI context. In the United
States of America, the number of animals per livestock operation has increased significantly

10
because large-scale facilities are more profitable, despite the fact that crowded animals are more
susceptible to disease, thereby increasing health costs (Tilman et al., 2002).
Livestock diet balancing and formulation is crucial to make best and most profitable use
of the feeds available in each unique production situation and deliver appropriate energy and
nutrients that allow animals to express their genetic potential for growth, development, and
production. It can also be important to minimize the excess of nutrients (those that will not be
absorbed and utilized by the animal) that would otherwise be excreted into the environment. This
practice is commonly known as precision feeding and has been defined as “feeding livestock so
that animal performance is not adversely affected but so that nutrient excretion to the environment
is the smallest quantity possible” (Cole, 2003). Other definitions including economic and social
aspects have also been suggested. Opportunities have been documented for N and P nutrition
(Cerosaletti et al., 2004; Vasconcelos et al., 2007) and feeding management (Vasconcelos et al.,
2006). Similarly, mitigation strategies for methane emission by livestock, especially ruminants,
have also been proposed (Eckard et al., 2010; Gerber et al., 2013; Knapp et al., 2014; Tedeschi et
al., 2011a; Tedeschi et al., 2003). Diet and feeding practices that have been reported (based on
survey analysis) to improve sustainability include: (1) minimize water pollution, deforestation, and
air pollution from an environmental perspective; (2) produce animal protein affordably without
competing with crop cultivation for human food or compromising ethical aspects of livestock
wellbeing; (3) reuse feed waste after ensuring its safety, and (4) provide incentives to those
adopting sustainable diet and ethical feeding practices (Makkar and Ankers, 2014). Many of these
goals, however, can only be achieved with the assistance of decision support systems through
computer modeling and simulation that accurately and precisely formulate diets that meet animal
demand for energy and nutrients for an optimized performance under various production scenarios.
Although mathematical models have been used to predict environmental impacts of
ruminant production (Tedeschi et al., 2014a) with varying results, under certain conditions they
can be used to explore areas of uncertainty of ruminant production on environmental impact and
Feed the Future issues. Furthermore, uncertainty caused by statistical variation can be
incorporated into stochastic models to forecast confidence regions. Forecasts are valuable to
quantify technology impact or alternative production options on environment and food production
as well as understand how these options will behave over time.
Mathematical models have expanded as our awareness of their potential for data mining
and processing becomes more apparent. Mathematical and computational models can assist in
addressing this deficiency through whole-farm modeling (WFM) simulation (Parsons et al., 2010;
Snow et al., 2014). Though animal WFM submodels can vary considerably among species or
systems (Tedeschi et al., 2014b), sustainability of beef (Rotz et al., 2013) and dairy (Rotz et al.,
2010) life cycles can be assessed through production simulations scenarios using WFM;
understanding how and when animals and crops affect the environment can provide timely
guidance for policymakers.
Data measurement and collection needed to understand livestock-environment feedback is
often very time consuming. Therefore, the development and deployment of technology to support
data collection for accurate and precise predictions will likely require a high degree of spatial
resolution (e.g., high-definition satellite imagery) as well as fast and reliable data acquisition tools
(e.g., unmanned aerial vehicles).
Sustainable production systems: Livestock and grazing systems
Invasive species are any alien animal(s) or plant(s) that disturb an ecosystem by displacing
more productive, nutritionally valuable native species and altering the normal behavior of natural
cycles (Peterson, 2003). Invasive species (e.g., feral species) resulting from climate change or
ecosystem disturbance can be a major hindrance to agriculture (Seward et al., 2004). They can be

11
useful, however, in developing resistant or resilient species or ecosystems. The genetic attributes
of invasive species that allow them to thrive in harsh environments can likewise ensure the success
of crops/grasses of interest in the same environmental conditions.
Droughts in the United States of America in 2011, northeastern Brazil in 2013, as well as
many other regions of the world, including southern Europe and most Mediterranean areas, raise
a concern over sustainability and resilience of both native and sown grasslands (Craine et al.,
2013). How can we identify herbaceous species that thrive under these stressful conditions?
Volaire et al. (2014) proposed a conceptual prototype to analyze adaptive responses of perennial
herbaceous species. They pointed out two major challenges for plant breeding and agro-ecology
research to cope with recurrent drought problems: 1) select new plant material that incorporates
long-term survival and persistence to droughts (i.e., drought resilience or drought resistant), and
2) use forage mixtures to ensure species diversity that would maintain a critical level of herbage
mass production through environmental fluctuations. Kahmen et al. (2005) pointed out the
importance of species diversity in grasslands not only for persistence but also for productivity,
especially below ground.
The use of agroforestry (e.g., agro-silvo-pastoral systems) is a potential SI approach for
grazing ruminants in several regions around the world. It is perceived that increased livestock
production could be achieved if smallholder mixed-crop-livestock systems, which are mostly
located in tropical regions, increase their productivity (Oosting et al., 2014). Campbell et al. (2014)
illustrated this approach through feeding cattle Leucaena leucocephala, a legume tree that is
widely available in tropical and subtropical ecozones. They reported that adding small amounts of
Leucaena leaves to dairy cattle diets can increase daily milk yield and weight gain by at least
threefold, besides decreasing methane produced per kg of meat and milk (Thornton and Herrero,
2010). In contrast, Rueda et al. (2003) indicated that even though supplementation with sorghum
grain increased milk production and growth by 25 to 50% per animal on cattle farms in Brazil’s
western Amazon, it was less profitable than current forage-only diets. In their case, SI could be
better achieved by judicious fertilization of grass-legume pastures and greater stocking density
rather than feed supplementation, highlighting the importance of mathematical model simulations
for making management decisions. Veysset et al. (2014) classified Charolais cattle farms into four
groups: specialized conventional livestock farms (100% grassland-based farms (GF), n = 7),
integrated conventional crop–livestock farms (specialized farms that only market animal products
but that grow cereal crops on-farm for animal feed, n = 31), mixed conventional crop–livestock
farms (farms that sell beef and cereal crops to market, n = 21), and organic farms (n = 7), and
analyzed their technical, economic, and environmental performances. The authors concluded that
although all types of farms had comparable economic performances, the mixed conventional crop–
livestock farms made heavier and consequently less efficient use of inputs, and had environmental
performances less than the specialized farms. The route to success of agroforestry (i.e.,
agroecology, mixed conventional crop-livestock) farms is not easy and it may have many
limitations for their long-term economic and environmental sustainability.
The next big problem: water scarcity
Available water for food production refers to the sum of green water (naturally infiltrated
into the soil) and blue water (water in rivers and aquifers). By 2050, Rockström et al. (2009)
estimate there will be a shortage of blue water for 59% of the world’s population while 36% of the
world population will face shortage of blue and green waters. These are seriously alarming
estimates.
Without taking into account the impact of climate change on agriculture, FAO (2011)
estimated that agricultural irrigation will have to increase 11% to meet consumer demand for high
quality food as the human population approaches 9 billion by 2050. This estimate could be

12
exacerbated by climate change (Schmidhuber and Tubiello, 2007). The forecast for agriculture is
grim: “climate change will significantly impact agriculture by increasing water demand, limiting
crop productivity and by reducing water availability in areas where irrigation is most needed or
has comparative advantage” (FAO, 2011). To make things worse, Hilker et al. (2014) indicated
that since 2000, precipitation has declined across 69% of the world’s tropical evergreen forest (5.4
million km2) and across 80% of all subtropical grasslands (3.3 million km2). The increase in
irrigation needs, the reduction in rainfall, and the decline in terrestrial water storage in some parts
of the world may indicate the beginning of tropical forest desiccation in the 21st century, leading
to negative cascading effects on global carbon and climate dynamics (Hilker et al., 2014).
Future Directions and Needs
Practical examples that can be used for SI in sub-Saharan Africa are available for livestock
(FAO, 2002) and crops (Juma et al., 2013). Nonetheless, some questions remain unanswered for
the sustainable livestock intensification concept. Will it adequately support the Feed the Future
initiative? What are the differences between regional and global application? Are rational
management and human intervention the keys to fostering resilient systems? The following are a
few examples among many.
Fetal programming
Maternal under- or over-nutrition during pregnancy are detrimental to growth,
development, and the function of some biological systems in offspring, including reproductive
development and efficiency (Mossa et al., 2015). Ferrell (1991) analyzed non-nutritional factors
that can alter fetal development through embryo transfer between large (Charolais) and small
(Brahman) genotype into recipient cows of similar and dissimilar genotype. Their data indicated
that at about 230 days of gestation, fetal genotype accounted for most of the fetal weight variation
among treatments, confirming that fetal genetics determine most of the fetal growth (Bell, 2005).
After 230 days of gestation, however, dam genetics became increasingly influential on fetal
weight, especially of Charolais fetuses transferred to Brahman dams. Fetal programming is based
on the developmental origins of health and disease (Gotoh, 2015) in which alteration in the
nutrition and endocrine status of the fetus or during the early postnatal phase can cause
developmental alterations that permanently modify structure, physiology, and metabolism of the
animal into adulthood.
Genetics and system sustainability
One of the basic principles of quantitative genetics is that, by changing allele frequencies,
the population mean for a given character can be changed. In the 20th century, livestock breeding
was very effective at implementation of selection improvement programs. The important traits of
the time were quantitative in nature (milk yield and growth rate) with moderate to large additive
genetic control in many populations supporting that selection. Extension programs in the United
States were important components of the overall improved agricultural environment of that
century, and technological advances, especially in the last half of the century, contributed strongly.
Although improvements in milk yield and beef quantity were not exclusively the result of
selection, no one will contest that the genetic component of this improvement was substantial. For
comparable amounts of milk yield, modern dairy production systems require only a fraction (one
third or less) of the animals, feedstuffs, water, and land as compared with systems in use in 1944
(Capper et al., 2009). For comparable beef production, modern systems require approximately 70,
81, 88, and 67% of the animals, feedstuffs, water, and land, respectively, that were required in
1977, while producing 82, 82, 88, and 83% of the manure, methane, carbon dioxide, and total
carbon footprint, respectively (Capper, 2011a). However, as growth increased for cattle in beef
production, mature BW also increased, accompanied by increases in daily resource use and GHG
emissions (Capper, 2013).

13
 As described by Capper and Bauman (2013), the “dilution of maintenance” concept is the
improvement of animal productivity such that a greater quantity (e.g., meat or milk) is produced
in a given time period. Systems with such economies of scale excel with respect to absolute
resource efficiency and utilization. Additionally there is an upper biological limit to traits like
growth and milk production (Capper and Bauman, 2013). The beef production system in the
United States of America is segmented; efficiency in one segment is often antagonistic to
efficiency in another segment. For example, although increased BW/growth rate of cattle
facilitates economies of scale from the point where cattle are fed high concentrates in feedlots
through slaughter and beef packaging, the heifer half-siblings of those steers are bred to join the
nation’s cow-calf herd. Body weight and other mature size measures of those females increase but
many aspects of reproduction are diminished with increased size (Vargas et al., 1999). Other
important components of the production system may be influenced negatively; for example,
greenhouse gases emissions such as methane would positively associate with mature cow size
(Capper, 2013). Smaller cows would require fewer resources for comparable production vis-á-vis
larger cows, even if they were equally productive from a fertility/maternal perspective. Although
more “traditional” extensive production systems may be perceived to be more sustainable from
many perspectives, the reality is often demonstrated to be otherwise (Capper, 2011b, 2012;
Stackhouse et al., 2011).
System sustainability should consider more than those genetic characters traditionally
representative of animal productivity (Hermansen and Kristensen, 2011; Van Eenennaam, 2013).
What appears to be missing in the evaluation of system sustainability is the inclusion of traits such
as cow reproduction that are considered less from an economy-of-scale perspective. The additive
genetic component of variation in such traits is low, making selection a more long-term process
with less visible results in the short term. This is the opposite of the 20th century success traits,
namely milk yield and growth or weight, which are certainly more responsive in the short term.
There are, however, genetic improvement approaches for such traits that are successful in the short
term, such as crossbreeding.
A second concern with current assessment of system sustainability could be the global
approach. Cow adaptation to local environments is a primary animal breeding strategy for
sustainable beef production (Olesen et al., 2000). For example, consider a resource-limited
environment, perhaps drought conditions. Cows can contribute to their population by 1) diverting
all their energy into procreation, often at the expense of their long-term reproductive efforts, or by
2) living another year but withholding nutrients and energy from the reproductive process. Our
own anecdotal related observations are that highly productive European breeds of cattle respond
with the former strategy and adapted, but otherwise less productive “indigenous” cattle breeds
often employ the latter. Population growth and demand for protein will grow most this century in
tropical developing nations (Laurance et al., 2014). Some aspects of adaptation to such
environments have been quantified and evaluated genetically (Riley et al., 2011; Riley et al., 2012),
but much potential characterization remains. This may not be as severe a concern if reproduction
traits are considered as an outward manifestation of adaptation.
Flexibility of response to environmental stress could strongly influence system
sustainability through environmental perturbations. Genetic flexibility appears to be an essential
part of livestock production system sustainability. Conservation genetics and diversity studies
illustrate one aspect of flexibility: diversity statistics indicating heterozygosity at markers and
genes throughout the genome (Van Eenennaam, 2013). Markers are visible but without function
(some researchers categorize markers into those without and with function), but believed to be
linked closely to unknown, unseen genes. Genomic markers may not be in coding regions
(genomic regions with some function, for example those coding for RNA that may or may not be
ultimately translated into proteins); coding regions may be more conserved, that is, they maintain
their identity without permitting alternative versions, because those may prevent protein

14
production. It would therefore be essential that diversity should not be considered exclusively from
a heterozygosity-of-marker perspective. Theoretically, continued selection for a single character
will reduce additive genetic variation; that is, it will tend to make animals homozygous for many
loci, reducing heterozygosity, and flexibility associated with having multiple alleles (alternative
versions of a gene). Meiosis as a diversity-producing mechanism would be offset by increased
homozygosity of individuals at many loci. As genes of importance continue to be characterized,
allelic diversity for those genes may be a worthy consideration in livestock planning and breeding
programs.
Preservation and conservation of breeds or races as they exist could maximize future
genetic flexibility as environmental perturbations are encountered. Breeds are populations with
allele frequencies that have been influenced by common (to the breed) forces across time. These
would be particularly important with respect to traits of adaptation to local conditions, as
redevelopment of these adaptations may be costly in terms of loss of individuals and of long-term
persistence of the population. This conservation may be a more prominent issue in developing
countries, as historically one pattern of improving productivity is to import productive breeds or
races and then attempt to alter the local environment to accommodate them. Capper and Cady
(2012) demonstrated differential breed contribution to a component of system sustainability
(environmental impact) in the cheese production.
It may be attractive to consider that genetic improvement could eventually result in highly
selected sets of animals that would excel in all situations. Of course, no animal is best at
everything; limited nutrient partitioning to growth, for example, depletes those nutrients available
for other life processes. Selection indices can be developed to assign weights to important
characters for selection. Although a global emphasis and perspective is beneficial overall (Herrero
and Thornton, 2013), improvement planning must be implemented at not too broad a geographic
scale, as different genotypes (aggregate) will perform unequally in local environments. An
objective would be to develop and promulgate systems of parental ranking (expected progeny
differences, predicted breeding values, or some other metric representing genetic merit) at the
smallest reasonable local level possible, at least at the country level. It would also seem to be
important to extend the scope of genetic investigation and implementation of advanced
methodology to smallholder livestock systems that are often characteristic of tropical protein
production. Those efforts at present are limited (Ejlertsen et al., 2012; Marshall et al., 2011).
Genomic improvement programs (Capper et al., 2008; Capper and Hayes, 2012;
Stackhouse et al., 2011) could have increasing impact as they develop as part of sustainability of
food production systems. Although this has not proceeded at a desired rate for selective
improvement of livestock, the expectation is that it will become reality at some point. As discussed
above, consumer perceptions and social acceptability of the technology will have an important
effect on designing sustainability systems (Croney et al., 2012; White and Capper, 2013);
sometimes the general public might be antagonistic to true sustainable systems and SI cannot be
achieved.
Conclusions
Sustainable intensification will be necessary to increase food production that meets the
needs of a growing world population. We will need to address multiple facets within animal
agriculture systems, especially in economically and environmentally challenged environments.
Feed efficiency is likely the single most important variable that can drive up output per input ratio
while concurrently reducing the carbon footprint of ruminant production by reducing greenhouse
gas emissions (i.e., methane) under climate-change conditions. Thus, the identification and
selection of feed efficient animals, feeding systems, and technologies that will improve efficiency
of nutrient use are priorities for sustainable livestock intensification programs. Taking full

15
advantage of genetic potential within animal genomes, whether from a breed or selection
perspective, will likewise contribute. Certainly investing these options within the world’s socially
and economically marginal populations that constitute today’s predominant animal agriculturalists
seems likely to have positive returns. Computerized mathematical models based on sound science
will be needed to integrate accumulated biological and management knowledge to identify and
implement the most productive, economically and environmentally sustainable system in each
unique production situation.
Pretty et al. (2011) drew several lessons from their experiences in spreading the use of SI,
including (1) synergize scientific knowledge and farmer experience into technologies and practices
that integrate crops and animal production with agroecological and agronomic management; (2)
develop social infrastructure that builds trust among individuals and agencies; (3) improve farmer
awareness, knowledge, and capacity through extension and other education programs; (4) engage
with industry and private partners to supply goods and consulting services; (5) focus on gender-
neutral educational, microfinance, rural banking, and agricultural technology needs; and (6) foster
public support for agriculture. Sustainable intensification is not the result of improved biological
and physical processes alone; it requires the intervention of human ingenuity to manage the system
appropriately and intelligently in an integrated way. The increase in food production has to be
achieved through enhanced yield (output/input) rather than expanding the area of land as the latter
may further increase environmental burden. Smallholder farmers may benefit the most from SI
that leads to greater integration into the market economy.
In deploying the technology, we concede that a catchy phrase or slogan such as sustainable
livestock intensification will not miraculously solve the problem at hand and it may inexorably
backfire if interest groups apply it in ways for which it was not intended, thereby corroding the
technology. Finally, the question that remains still unanswered is: will SLI effectively increase the
supply of animal products for those who would benefit the most? From a broader perspective, a
more daunting question is whether global agriculture in 35 years will be able to feed 10 billion
people, produce biofuel and clean electricity, supply fiber, and at the same time reduce its
environmental footprint to avert or even reverse climate change?
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