Animal Feed Science and Technology 166167 (2011) 291301

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Animal Feed Science and Technology

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Lowering ruminant methane emissions through improved feed

conversion efciency
G.C. Waghorn a, , R.S. Hegarty b
a
DairyNZ, cnr Ruakura & Morrinsville Roads, Hamilton 3240, New Zealand
b
Beef Industry Centre of Excellence, Industry and Investment NSW, Travenna Road, Armidale 2351, NSW, Australia

a r t i c l e i n f o a b s t r a c t

Improvements in feed conversion efciency (FCE) can be applied to individual animals as

well as to production from land, as in a farm system. Our focus relates mainly to food pro-
duction from individual animals within any animal population where there is divergence
Keywords: in the efciency that individuals use ingested feed for maintenance and production; pri-
Feed efciency marily due to differences in digestion and metabolism. Intake variation from the predicted
Methane mean for individuals of a similar size and level of production in a population has been
Residual feed intake
termed residual feed intake (RFI), with low values indicating an efcient animal. Efcient
Emissions intensity
animals require less feed than average and can be expected to produce less CH4 and N2 O
RFI
per unit product than the population average at a similar level of production. Selection for
this trait will lower CH4 emissions per animal, unless more animals are kept to eat the feed
not required by efcient animals. There are few published evaluations of CH4 yields from
animals with divergent RFI and there is little evidence that efcient animals have a differ-
ent CH4 yield expressed as CH4 /kg dry matter (DM) intake. Of equal or greater importance
than RFI is the need to select high producing animals, as this will reduce emissions/unit of
product, referred to as emissions intensity (Ei). Research should identify productive indi-
viduals that have a low RFI to minimise Ei and maintain food production. The extent to
which CH4 can be reduced by selection for RFI will depend on the heritability of efciency,
dispersal of efcient animals through all populations and their resilience in a production
system (i.e., robustness). The benet of RFI to lowering greenhouse gas (GHG) emissions is
its application, irrespective of farming system (i.e., conned, intensive, extensive grazing),
especially because efcient animals are likely to increase farm protability. Efcient ani-
mals are already in all herds and ocks and research must identify and remove inefcient
individuals, while retaining and ensuring efcient ones are t to purpose. However, the
biggest benets to reducing emissions and increasing production will be associated with
good animal management practice (e.g., appropriate genetics, reproductive performance,
longevity) with efcient animals superimposed. Good animal systems management will
improve protability, and apply to both intensive and extensive systems to increase food
production and lower Ei. One dilemma for agriculturists will be the practice of feeding
grains to ruminants, as gains in animal efciency, especially in reduction of Ei, are likely to

Abbreviations: Ei, emissions intensity (g/product); FCE, feed conversion efciency; GHG, greenhouse gas; LW, liveweight; ME, metabolizable energy;
MY, methane yield; OM, organic matter; RFI, residual feed intake.
Corresponding author. Tel.: +64 7 858 3881; fax: +64 7 858 3571.
E-mail address: Garry.waghorn@dairynz.co.nz (G.C. Waghorn).

0377-8401/$ see front matter 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.anifeedsci.2011.04.019
292 G.C. Waghorn, R.S. Hegarty / Animal Feed Science and Technology 166167 (2011) 291301

be biggest with high energy density rations, but feeding grain to ruminants may become
an unsustainable practice if food supplies for humans are limited.
This paper is part of the special issue entitled: Greenhouse Gases in Animal Agriculture Finding
a Balance between Food and Emissions, Guest Edited by T.A. McAllister, Section Guest Editors:
K.A. Beauchemin, X. Hao, S. McGinn and Editor for Animal Feed Science and Technology, P.H.
Robinson.
2011 Elsevier B.V. All rights reserved.

1. Introduction

Efciency of feed utilisation can be described by a number of metrics which combine measures of feed consumption
and desired animal production. Of these, residual feed intake (RFI), is being widely adopted for genetic improvement of
ruminants (Archer et al., 1999). Recognition that feed consumption also generates an undesirable animal product (i.e., enteric
CH4 ) has seen efciency of conversion of feed into enteric CH4 (or CH4 yield; MY) emerge as an important consideration
and potential animal and plant breeding objective (Molano and Clark, 2008). As the long term viability of animal production
may be dependent upon the balance of desirable and undesirable outputs, the term emission intensity (Ei; CH4 /unit animal
product) has been created (Leslie et al., 2008). The interplay of the indices RFI, MY and Ei is shown in Fig. 1. While the metrics
of RFI, MY and Ei are typically calculated at the individual animal level, they can equally be applied at the whole farm level.
This review will interpret opportunities for inuencing these outputs and indices at both the individual animal and farm
system level through changes in feed efciency using RFI as the metric of choice.

2. Feed conversion efciency in ruminant systems

Extensive grazing systems have fewer inputs, lower costs and fewer options for greenhouse gas (GHG) mitigation than
intensive animal production systems (Eckard et al., 2010). However, management of feed and animal genetics bases are
potential points of intervention for affecting production and protability, as well as to reduce Ei from extensive (Bentley
et al., 2008) and intensive (Donoghue et al., 2009) ruminant livestock systems. Changes in efciency of feed utilisation can
improve production and alter CH4 emissions.
Feed accounts for 6677% of costs in beef cow/calf and feedlot nishing systems in North America (Williams and Jenkins,
2006) and about 50% of production costs in New Zealand pastoral dairying systems (DairyBase, 2009; Beever and Doyle, 2007).
In grazing systems, efcient feed use is a function of harvest of available DM by animals (Orr et al., 2010) and efciency of
conversion of ingested feed to product.
Animal and farm management practices that affect feed use efciency include multiple aspects of animal genotype,
available feed and animal health, and reproductive management (Table 1). While Table 1 is not exhaustive, these variables
are important because they impact feed use efciency, as well as protability, which is the principal driver of ruminant
agriculture in the developed world.
Food creation as animal products is commonly reported as weight of feed consumed per weight of animal product
(e.g., liveweight (LW), carcass weight, milk production) generated (i.e., feed conversion ratio) or its inverse product/unit
feed which is referred to as gross efciency. While these are simple to calculate, they are not suitable as traits for genetic
improvement of feed use efciency in ruminants as they include correlated changes in intake and growth (Arthur et al.,
2001a,b). For this reason, RFI is the preferred metric for genetic improvement in feed use efciency.
It is essential that selection for low RFI animals has no adverse effect on animal productivity. Research has not demon-
strated important negative associations with RFI, but it will be important to monitor economically relevant traits. For
example, cattle with superior breeding values for RFI (i.e., efcient individuals) have less body fat, especially rib fat (Herd and
Arthur, 2009; Egarr et al., 2009), and implications of this for post-partum anoestrus in the grazing herd could be negative.
The risks of selection for one trait, such as milk yield, while ignoring others such as fertility, are well known, especially

Dry Matter
Intake

RFI MY

Animal Methane
Production Production
Ei
Fig. 1. Linkages between feed consumption and the output of desirable and undesirable (methane) animal products through residual feed intake (RFI),
methane yield (MY) and emission intensity (Ei).
 G.C. Waghorn, R.S. Hegarty / Animal Feed Science and Technology 166167 (2011) 291301 293

Table 1
Animal and key farming system variables that affect feed conversion efciency of ruminants and consequently the emissions intensity of the farming
system.

Animal options
Species sheep beef, dairy
Stage of maturity affects energy partitioning
Genetic potential (production, RFI) selection for protable traits
Environmental adaptation productive and reproductive resilience
Good grazers; seek out feed and water, even if substantial walking is needed. Low susceptibility to disease (ticks, internal parasites, etc.), avoid toxic
forages
Farm system options
Extensive grazing options often dictated by needs for sustainability and need to accommodate adverse conditions (feed supply) with minimal inputs
Provision of pastures or browse able to match animal energy and nitrogen requirements
Use of tested sires at regionally appropriate male:female ratio
Pregnancy scanning and removal of non-pregnant females
Seasonal supplementation of females to ensure return to service and of progeny to minimise time to nish or mating
Manage for minimal parasite impact.
Good mothering and high weaning rate; good growth rates
Intensive grazing use of improved forages, ready access to livestock, ability to supplement and manage to achieve a high production/ha
Production affected by forage type and grazing management to maximise pasture harvest
Optimal forage species to maximise feeding value
Use of additives, rumen modiers and supplements to ensure adequate feeding at all times
High reproduction, early mating, high conception and weaning rates
Attendance to health, good longevity to minimise replacement rate
Select animals that are adapted and have good genetic merit under forage grazing
Intensive feedlot for (dairy) production or nishing for carcass production
Optimal ration formulation and adaptation to nishing ration
Appropriate parasite and disease control on induction
Aware of heat stress risk & shade/sprinklers provided as appropriate

in the dairy industry (Wall et al., 2007; McDougall, 2006), and care must be taken to monitor all aspects of animal health,
reproduction and production in selection.

2.1. Residual feed intake (RFI)

Efcient animals (i.e., low or negative RFI) eat less than the average predicted based on dened LW, physiological state
and production, whereas inefcient (i.e., high RFI) individuals eat more than the predicted average. Selection for divergence
in RFI identies individuals which differ in feed requirements, but is independent of LW or productivity. Selection based on
RFI has been exploited in genetic improvement programmes with non-ruminant species for decades and has resulted in fast
growing animals requiring less feed (Hermesch, 2004). RFI is moderately heritable in beef animals (h2 = 0.140.46, Arthur
et al., 2001a,b; Pitchford, 2004; Arthur and Herd, 2005; Basarab et al., 2005; Lancaster et al., 2009) with differences in DM
intake of 15% after two generations of selection (Herd et al., 2002a). Testing for RFI is becoming more common in the North
American beef industry where grain and silage based diets are fed. In contrast to fresh pasture, these diets are higher in DM
and energy content, less bulky, do not deteriorate appreciably when fed once or twice daily, and have a consistent chemical
composition over the entire feeding period. While the RFI trait is conserved in grazing animals (Herd et al., 2002a), more
data are needed to correlate RFI of cattle measured with grain and silage based diets with RFI of cattle grazing pasture.
Current protocols for RFI determination in cattle require individual animal identication and accurate measurement of
intake as well as production (i.e., LW gain). Under intensive systems, feed is placed in bins on load cells with continuous data
capture, and animals feeding from each bin are identied by electronic identication. The cost of determining RFI is primarily
that of the equipment required and data interrogation. However measurement of RFI from cattle that are on pasture incurs
substantial additional costs because of investment in equipment and provision of suitable forage. This may be green chop
or dried forage that enables more accurate measures of DM intake, and costs are compounded by the duration of intake
measurements (e.g., 70 d for beef cattle; Archer et al., 1997). However, increasing weighing frequency from 714 d to 23 d
can reduce the measurement period to about 50 d (G.C. Waghorn, DairyNZ, Hamilton, New Zealand; unpublished data) and
use of week-on/week-off data collection would double the number of animals measured per period (Donoghue et al., 2009).
High measurement costs have also prompted searches for genetic markers to identify efcient individuals more easily (Chen
et al., 2009).
Current research is extending measurements of RFI to dairy cows fed forages (Carnie et al., 2010). In that study, RFI for
milk production was determined by measuring RFI for daily gain in pre-pubertal dairy heifers fed alfalfa cubes. Use of daily
gain as a proxy for lactation efciency is based on two assumptions, being that divergence will apply to metabolizable energy
(ME) requirements for maintenance, which accounts for more than half a dairy cows lifetime feed needs (Freer et al., 2007),
and that RFI has a physiological/biochemical basis so that efciency for gain will also apply to milk synthesis. Once selected,
efcient individuals require less feed for the same level of production (Table 2) and this will lower Ei (Table 3). At the farm
level, a consequence of lower feed requirement per individual may be to increase stocking rate (i.e., animals/hectare). Thus,
under this scenario, total GHG would not decline, although there would be more production from the same land area.
294 G.C. Waghorn, R.S. Hegarty / Animal Feed Science and Technology 166167 (2011) 291301

Table 2
Examples of differences in feed intake (kg DM/d) of cattle identied with divergent residual feed intake (RFI) at similar levels of productivity.

Animal age, type and average daily gain (ADG)a Low RFI (Efcient) High RFI (Inefcient) Sourceb

Steers, 78 months; ADG 1.5 kg/d 8.00 8.93 1

Cows at maintenance 13.1 16.5 2
Progeny of divergent cows; ADG 1.3 kg/d 8.3 9.1 2
Bulls, ADG 1.75 kg/d 9.60 12.00 3
Steers, ADG 0.97 kg/d 7.27 10.13 4
Calves, ADG 1.06 kg/d 8.73 10.86 4
Calves, nishing 1.42 kg/d 8.45 10.63 4
Steers, nishing 1.48 kg/d 11.7 12.7 5
Steers, nishing 1.12 and 1.22 kg/dc 8.38 14.13 6
Steers, nishing 1.48 and 1.46 kg/d 9.62 11.62 7
Steers, nishing 1.11 and 1.07 kg/d 10.4 11.1 8
Bulls, ADG 1.42 and 1.40 kg/d 8.62 10.28 9
a
A single value for average daily gain (ADG) indicates a similar value for low and high RFI animals, two values correspond to ADG for low and high RFI
respectively.
b
1: Basarab et al. (2003); 2: Basarab (2005); 3: Fox et al. (2004); 4: Carstens and Tedeschi (2006); 5: Hegarty et al. (2005); 6: Hegarty et al. (2007); 7:
Nkrumah et al. (2006); 8: Herd et al. (2009); 9: Lancaster et al. (2009).
c
Animals used for CH4 measurement, summarised in Table 4.

It is important that animals selected for low RFI (i.e., requiring less feed than predicted) have been selected for high
productivity, whether this be as daily gain or milk production. While RFI is independent of production, high producing
animals are the most protable because they create more product and maintenance energy costs are diluted relative to
lower producers (Table 3). For this reason, Arthur and Herd (2005) proposed a 2 stage selection, to identify animals with a
high genetic merit, with only the top 1020% most productive bulls being tested for RFI.

2.1.1. Physiology of RFI

The bases for differences in RFI have been ascribed to the physiology and biochemistry associated with digestion, energy
capture (i.e., as ATP) and energy utilisation, but these have not been studied in detail. In their analyses of Angus steers
selected for divergent RFI, Herd and Arthur (2009) examined feed intake, feed digestion, metabolism, physical activity and
thermal regulation. They were able to explain 73% of the variation in RFI, and attributed it to variation in protein turnover,
tissue metabolism and stress (37%), with lesser contributions from digestibility (10%), heat increment and fermentation
(9%), physical activity (9%) and body composition (5%). One reason advanced by Basarab et al. (2003) for lower energy
requirements of efcient steers was the smaller visceral mass (i.e., stomach, intestines, liver) in animals with low RFI (51.1 kg
versus 55.3 kg for inefcient steers). Although a smaller mass would lower energy expenditure, it is hard to reconcile with
higher digestibility. Higher digestibility can result from a longer rumen retention time and a larger digesta pool, traits that
are typically associated with a larger rumen. Recent measurements from divergent Angus-Hereford steers by Cruz et al.
(2010) did not identify differences in visceral mass.
Differences in digestibility among animals fed the same diet offers potential for creating divergence in RFI because of a
higher nutrient energy yield, but more digestion is likely to increase CH4 yields as more H2 derived from feed will be used to
create CH4 . Although Richardson and Herd (2004) suggested a divergence of only 2% in DM digestion (i.e., higher in efcient
steers), Nkrumah et al. (2006) reported means of 75.3%, 73.4% and 70.9% for high, medium and low feed efcient steers.
In his review of 80 studies, Titgemeyer (1997) reported variation among individuals within experiments of 1.217.4% in
rumen digestion and 0.49.1% for total tract digestibility. Thus there appears opportunity to affect nutrient availability and
methanogenesis by selecting for efciency of digestion.
Variation in rumen function among individuals fed the same diet was reviewed in relation to RFI in cattle by Waghorn and
Dewhurst (2007). There is evidence that individual animals harbour their own specic microora, as demonstrated by the
heritability of susceptibility to legume bloat (Morris et al., 1991), and differences in rates of ruminal in sacco feed degradation
among animals fed the same diet (Weimer et al., 1999). Variation may include the extent and duration of digestion in the
rumen (i.e., residence time), which appears to be a heritable trait in sheep (Smuts et al., 1995), which inuences quantities

Table 3
An illustration of changes in emissions intensity (Ei; CH4 /kg live weight gain) in sheep fed diets of varying quality.

Dietary ME (MJ/kg DM) Forage Gain (g/d) Methane (g/kg DM intake) Feed:gain ratio (kg DM intake/kg gain) Ei (g/kg gain)

10.0 Ryegrass pasture 100 24.0 13.6 300

11.0 Ryegrass pasture 150 22.0 9.4 210
12.0 Ryegrass pasture 200 21.0 7.5 160
11.5 Lucerne 250 20.0 6.7 130
12.0 Sullaa 300 17.5 6.2 110

ME: metabolisable energy; DM: dry matter.

Calculated CH4 emissions per unit of liveweight gain from growing lambs fed forages with a range of feeding values (from Waghorn and Clark, 2006).
a
Sulla (Hedysarum coronarium) contains condensed tannins.
 G.C. Waghorn, R.S. Hegarty / Animal Feed Science and Technology 166167 (2011) 291301 295

Table 4
Feed dry matter (DM) intakes and CH4 production from cattle selected for variance in residual feed intake (RFI). Low RFI are efcient, high RFI are inefcient.

Number of cattle Dry matter intakes Production (kg/d) CH4 production (g/d) g CH4 /kg DM intake Source

RFI Low High P Low High P Low High P Low High P

Steers 10 + 10 8.38 14.13 0.001 1.13 1.23 ns 131 173 0.09 16.3 14.7 0.37 1
Steers 11 + 8 8.65 8.68 0.39 1.48 1.46 97.5 129.3 0.04 11.3 14.9 0.04 2
Cowsa 8+8 10.8 11.2 0.57 na na 191.8 193.9 0.78 18.3 17.5 0.40 3
Cowsb 8+8 12.2 12.5 0.61 na na 272.1 258.7 0.39 22.4 20.9 0.21 3

References and methodology:

1: Hegarty et al. (2007); 590 kg steers fed diets containing 750 g/kg grain (DM basis).
2: Nkrumah et al. (2006), 500 kg steers fed diets containing 80% grain (DM basis) with CH4 measured 16 h/d without access to feed, Daily gain was from
previous feedlot trials where intakes were 9.6 and 11.6 kg DM/d for efcient and inefcient animals.
3: Waghorn et al., DairyNZ, Hamilton, New Zealand; unpublished data.
a
Nonlactating Holstein/Friesian aged 21 months fed alfalfa cubes.
b
Holstein/Friesian aged 26 months at day 60 of lactation fed pasture.

and proportions of volatile fatty acids, proteolysis and deamination of amino acids, microbial growth and methanogenesis
(Janssen, 2010). Differences in products of digestion and rumen outow affect H2 availability for methanogenesis (Janssen,
2010), and energy capture efciency as high energy phosphate bonds by ruminal microorganisms (Waghorn and Dewhurst,
2007).

2.1.2. Gains in productivity through RFI

Differences in DM intake and productivity of beef cattle with divergent RFI are indicated by examples in Table 2. Dif-
ferences between selections are substantial, with the magnitude of the differences dependant on the proportion of the
screened animals selected, with differences between the most and least efcient individuals biggest when they comprise
5% as opposed to 20% of the population. Furthermore, the extent to which feed intakes can be reduced by selection is nite,
because energy must be allocated to maintenance and synthesis of meat and milk. Selection for RFI is independent of prod-
uct composition, and so the amount of energy per unit product cannot be altered. The extent to which individual intake is
reduced over generations depends on the heritability of the trait and the selection pressure applied. Mitigation of livestock
GHG emissions will be affected by the rate and extent to which the trait can be disseminated throughout a population. Bene-
ts of selecting sires as opposed to dams for improving RFI are clear, especially when articial insemination is in widespread
use.
However, it is important to appreciate that efciency gains in a herd or ock from selecting efcient animals will be
about half of that indicated by the breeding value for RFI, unless retention of efcient animals is accompanied by removal of
inefcient individuals. The average feed intake by individuals in a population comprises that from animals with all levels of
efciency, and so improvement is best achieved by animal culling as well as retention. Furthermore, gains in progeny feed
efciency will be one half of that indicated by the RFI breeding values of the parents.
In general, the impact of selection for RFI on dairy cattle will be less than for beef cattle because a higher proportion of
feed energy will be expressed in milk than is retained in LW gain. The energy in a product cannot be affected by variation in
RFI (i.e., conservation of energy), and product composition is largely independent of RFI (Arthur et al., 2001a,b; Richardson
and Herd, 2004; Lancaster et al., 2009). The energy content of milk and LW gain vary in response to proportions of fat, protein
and water (e.g., affected by diet, stage of lactation, age of animal) but these differences will be similar for low and high RFI
selections. Typical values for 1 kg empty LW gain in growing cattle are 1924 MJ/kg, representing about 22% of ME intake
(Freer et al., 2007). In contrast, the energy in milk of cows producing 2030 kg/d represents 3541% of their respective ME
intakes (Freer et al., 2007). Only the energy used for product synthesis and animal maintenance is able to be manipulated
through selection, but this example suggests that about 78% of ME is amenable to selection in growing cattle, compared to
5965% of ME in lactating cows.

3. Methanogenesis from animals selected for divergent RFI

Three studies where CH4 production was measured from animals selected for divergent RFI are summarised in Table 4.
Measurements made with steers have used either high release rate of SF6 over 5 d (Hegarty et al., 2007), or 16 h measurements
in respiration hoods repeated at 3 d intervals (Nkrumah et al., 2006). More recent measurements from cows have been made
over 48 h in respiration calorimeters (G.C. Waghorn, DairyNZ, Hamilton, New Zealand; unpublished data). Values from these
studies (Table 4), varied from about 11 to 22 g CH4 /kg DM intake with data being derived from concentrate based diets
containing 7580% grain fed to steers as well as alfalfa cubes and fresh ryegrass pasture fed to dairy heifers. This data
should be interpreted with care, especially where feed intake was restricted during CH4 measurements, because emissions
may be twice as high during and immediately after eating, than at other times (G.C. Waghorn, DairyNZ, Hamilton, New
Zealand; unpublished data). The CH4 yield was substantially lower from cattle fed concentrate diets over 16 h (Nkrumah
et al., 2006) and, although Hegarty et al. (2007) showed a relationship between CH4 production (MP, g/d) and RFI (kg/d):
MP (g/d) = 13.3 RFI + 179.5; P=0.002, R2 = 0.12, there were differences in CH4 yield among animals selected for varying RFI
296 G.C. Waghorn, R.S. Hegarty / Animal Feed Science and Technology 166167 (2011) 291301

Table 5
Emissions, expressed as CO2 -e, associated with production of foods for human consumption in the United Kingdom.

Food Type of emissions (g/kg product) Emissions expressed as CO2 -e (kg)

CH4 /NH3 /N2 O /kg product /MJ edible product /kg edible protein

Milk 19/3/1 1.0 0.37 28.6

Eggs 8/28/4 3.8 0.57 33.3
Chicken meat 5/23/3 3.5 0.40 18.4
Pork 49/28/2 4.7 0.30 34.2
Beef 265/71/12 14.7 1.40 93.5
Sheep meat 301/41/11 15.8 1.51 92.9
Wheata 0.20.8b 0.010.04 0.10.4
Peasa 0.1 0.005 0.04

From DEFRA (2008) and Gill et al. (2010).

a
Derived from Canadian research by Khakbazan et al. (2009) and excludes soil carbon losses from cultivation.
b
Range is due to variations in inputs and grain yield.

(Table 4). Daily CH4 production from cattle with low RFI (i.e., efcient animals) likely declines as DM intake is reduced, but
more information is needed to conrm effects of RFI on CH4 yield from cattle fed high concentrate diets. Cattle of divergent
RFI fed equal DM intake of a high concentrate diet did not differ in CH4 production (Hegarty and Herd, 2008).
Selection for low RFI should result in ruminants which produce less CH4 at similar levels of production, so that their Ei
should be lower (Tables 3 and 5) as less feed will be required for production. Reductions in emissions and Ei with improved
RFI should also apply to N2 O (Herd et al., 2002b), as lower N intakes will result in less urinary N excretion and reduce
N excretion/unit production. Consequently, selection for RFI is benecial for producers, consumers and the environment
(Beukes et al., 2010) because more food is produced with less GHG emissions and more prot. RFI can be applied to both
intensive and extensive systems and to all animals used for food production.

4. Emissions intensity and food production

Ruminant production systems have a high Ei relative to pork and poultry production systems (Table 5), largely because
of enteric CH4 generation (DEFRA, 2008). A low Ei or greenhouse cost per unit of animal product is increasingly regarded as
a sustainability criterion by livestock producers and consumers, and the value of ruminants for their contribution to global
food supply and GHG is under scrutiny. There are two main issues concerning use of Ei to manage ruminant production and
prioritise feeds used for food production, being the GHG emissions associated with production of edible food (milk, meat)
and the amount of human edible feed that is fed to ruminants (mainly grains) to create ruminant products.
A simple summary (Table 3) relating CH4 production and Ei to sheep growth demonstrates the importance of rate of gain,
forage quality and dilution of maintenance costs to lowering Ei. However, evaluation of farming systems based on life cycle
analyses, summarised by Beauchemin et al. (2010), indicated the level of complexity and range of assumptions required
to calculate, monitor and estimate emissions in complex farming systems. In their analysis of Canadian beef production,
Beauchemin et al. (2010) attributed 63% of total GHG emissions to enteric CH4 production, with N2 O accounting for 27%.
These and other life cycle analyses highlight the importance of factors such as longevity and reproductive efciency to dilute
GHG emissions associated with maintenance of the dam, as well as the need for rapid growth (Table 3) and low RFI to
minimise emissions from offspring grown for slaughter (Alford et al., 2006; Alcock and Hegarty, 2011).
Emissions of GHG from ruminants are in Table 5, with values for pork, chicken and other meats. The lower emissions
from non-ruminants are evident and, although life cycle analyses are central to calculation of Ei, as the assumptions and
inputs play a major role in determining outcomes, especially when comparing systems. Gill et al. (2010) report a doubling
of the calculated livestock emissions from 9% of global anthropogenic GHG to about 18% when all input costs, including land
use change and food processing, are considered. As a consequence, there are wide variations in predictions of Ei associated
with the same level of milk production (i.e., 0.51.5 kg CO2 -e/kg milk (Martin et al., 2010)). Similar variation is evident for
beef production, with a range from 17 to 37 kg CO2 -e/kg carcass, or a mean of about 33 kg CO2 -e/kg edible cuts (Beauchemin
et al., 2010).
Use of Ei enables a balance to be achieved between demand for food and the GHG costs associated with its production,
but analyses nearly always ignores losses of soil C associated with cultivation unless there is a change of land use. These
losses, often 10002000 kg C ha/yr, represent a depletion of soil organic matter (OM), but are lessened with minimal tillage
systems (Reicosky et al., 1995). However, depletion of soil OM in many cultivated soils demonstrates an overall loss of soil
C to the atmosphere when grains are grown. These losses are only taken into account in inventory if there is a change in
land use, and they are often ignored in life cycle analyses. Indeed the high productivity and lower CH4 yield associated with
high grain diets (>80% of DM intake) fed to ruminants (e.g., Beauchemin et al., 2010) suggests grain feeding could be a way
to reduce Ei relative to forages but, in view of C losses from cultivated soils, this premise should be challenged with new
research.
A detailed analysis of a wheat/pea cropping system in Canada by Khakbazan et al. (2009), showed GHG emissions asso-
ciated with wheat production (average annual yield 2500 kg/ha) ranged from 0.2 to 0.8 kg CO2 -e/kg. Emissions from peas
 G.C. Waghorn, R.S. Hegarty / Animal Feed Science and Technology 166167 (2011) 291301 297

(0.1 kg CO2 -e/ha) were much lower because less fertilizer N was used, but neither of these analyses took losses of soil OM into
account, or other GHG emissions associated with machinery manufacture and use. Although soil OM losses from cultivation
and cropping will vary widely with soil type, use of no till cropping may lower annual average losses (Reicosky et al., 1995;
Maljanen et al., 2007) to only 1000 kg/ha, resulting in emissions of about 0.4 kg CO2 -e/kg wheat. If these values were applied
to barley grain fed to fattening beef (Beauchemin et al., 2010), emissions would increase by 1.52 kg CO2 -e/kg carcass.
Losses of soil OM are well documented and usually detrimental to soil quality and crop production, but OM can be replen-
ished and there is provision in the Kyoto protocol for soil C sinks to be used to offset GHG emissions from grazing. Farming
systems evolve to optimise protability, and some environments are better suited to cropping than grazing, especially where
over-grazing could destroy an ecosystem. GHG emissions are only one aspect of the balance between food production and
environmental conservation and, although it may appear unreasonable to attribute a lower CH4 yield from enteric digestion
to ruminants fed high grain diets without considering all production factors, an alternative approach would be to reward
operations for producing ruminant products from pasture systems. Grazing does not require machinery to cut, dry, trans-
port and feed animals, but the best evaluation of ruminant systems should be based on a life cycle analysis incorporating all
inputs.
With regard to feeding human edible foods to ruminants, Gill et al. (2010) have calculated efciencies of systems used
for providing human edible animal products (i.e., meat and milk), and showed the energetic efciency of converting animal
feed (i.e., forages and concentrates) into milk, beef, pork and poultry meat was about 0.25, 0.06, 0.21 and 0.20, respectively.
Further assessment of the efciency of dietary protein use for protein production in these same products were estimated at
0.20, 0.07, 0.17 and 0.33, respectively. Efciencies were always lowest for beef production, but capture of energy in milk was
similar to that for pork and poultry production. These data were based on systems in the USA and South Korea, with values
being similar between countries. However, when the calculations assessed livestock production systems based on their
ability to convert human inedible feeds into to human edible products, ruminants were 23-fold more efcient than pigs
and poultry in the USA (Table 5) and 10 times more efcient than pigs and poultry in South Korea. The difference between
countries was a consequence of lower grain inputs to ruminant diets in South Korea. Presently, there are no incentives to
separate components of ruminant diets into human edible and inedible fractions, but if global demand for food begins to
exceed supply, ruminants would retain their value by producing high quality foods from human inedible forages, albeit with
a higher Ei than current values.

5. CH4 yield variation and measurement

In any set of CH4 production measurements, individuals vary in their CH4 yield (Blaxter and Clapperton, 1965; Sauvant
and Giger-Reverdin, 2009; Yan et al., 2010). Variation indicates potential opportunities for animal selection, provided the
differences are real, persistent and heritable. It appears that differences among some individuals in CH4 yield persist over
time (Robertson and Waghorn, 2002; Pinares-Patino et al., 2003; Vlaming et al., 2008) and understanding these differences
could provide further opportunities to mitigate impacts of ruminant animal production systems on GHG emissions. However,
some variation in CH4 yield has been attributed to the technique used for CH4 measurement, especially the SF6 technique
(Pinares-Patino et al., 2008), leading to the conclusion that the extent of variation among individuals may be less than
previously reported (Hammond et al., 2009). Thus, more research is needed to quantify individual animal variance in CH4
yield using accurate methodology to enable heritability to be estimated in a manner that could be employed for animal
selection to lower GHG emissions.
Components of rumen function affecting CH4 yield include level of intake, diet composition, products produced in the
rumen and the extent of digestion and possible anti-methanogenic comounds in the diet such as condensed tannins (Grainger
et al., 2009). Most factors are interrelated. For example, high feed intakes will result in high CH4 production, but the reduced
rumen residence time associated with high intake will tend to lower CH4 yield per unit DM intake (Hegarty, 2004). The decline
in CH4 yield with increasing intake above ME maintenance applies to both fresh pasture (Muetzel et al., 2009) and concentrate
diets (Yan et al., 2010). In a meta-analysis of several hundred measurements derived from calorimetry experiments, Sauvant
and Giger-Reverdin (2009) showed the decline in CH4 (g/kg digestible OM (DOM) intake) for multiples of ME maintenance
intake (M) to be: CH4 = 33.4 4.10 M. Average values were 26 5.5 g/kg DOM and average OM digestibility was 66 7.5%.
Methane yield with increasing proportions of concentrate in the DM decreased curvilinearly with effects being biggest
when intakes exceeded twice ME maintenance requirements. These ndings, based on data from sheep, cattle and goats,
support conclusions of Blaxter and Clapperton (1965) that decreases in yield with increasing intakes were largest with highly
digestible diets, and suggest the reductions in CH4 yield from selection of efcient animals (i.e., low RFI) will be biggest when
they have high intakes and are most productive.
Possibly the biggest challenge in measuring CH4 yield in individuals is to ensure that data are valid, which requires accurate
measurement of CH4 production and feed intake. If either CH4 or intakes are measured indirectly (e.g., using markers) they
may not be sufciently precise to distinguish small differences among individuals. For example, the SF6 technique has
been shown to account for substantial variation among animals, although the mean values for CH4 yield calculated using
this technique (Clark et al., 2006) are similar to those calculated by calorimetry (Hammond et al., 2009). There is a small
signicant correlation between rate of SF6 marker release and CH4 production (Pinares-Patino and Clark, 2008), but SF6
does not measure losses of CH4 via atus. Comparisons of CH4 yield measured by SF6 and calorimetry suggest SF6 values are
about 8% lower than chamber values (Grainger et al., 2007).
298 G.C. Waghorn, R.S. Hegarty / Animal Feed Science and Technology 166167 (2011) 291301

Additional concerns about the SF6 technique arise from comparisons of CH4 yield from sheep fed ryegrass with those
fed either white clover or chicory (Chicorium intybus). Previous measurements using SF6 (Waghorn and Woodward, 2006),
showing low CH4 yield from sheep fed chicory (16.2 g/kg DM) or white clover (1216 g/kg DM) were not conrmed by
calorimetry. Both Sun et al. (2011) and Hammond et al. (2011) have reported similar CH4 yield from fresh ryegrass, white
clover and chicory (22 g CH4 /kg DM intake), meaning that the accuracy of the SF6 technique may be affected by feed type.
However, the similar CH4 yield from these three forages is logical, because similar digestion of them suggest similar H2
release, and consequently CH4 production.
Accurate estimates of feed intake of ruminants conned to pens can be relatively easily obtained using electronic feed
intake systems, whereas use of markers (e.g., alkanes, Cr, indigestible bre) to measure forage intakes of individual grazing
animals is generally inaccurate. In theory, the alkane technique is an ideal method for determining intakes of individuals,
but with pastures containing many forage species, variability in the concentrations of alkanes in forage consumed and
poor faecal recovery renders this technique unreliable. Use of indigestible markers to determine intakes of grazing animals
assumes a similar digestibility of forage among individuals, an assumption that is unlikely to be true, and may be responsible
for some of the divergence in RFI. In some reports of CH4 yield, animal intakes have been based on values from feeding tables,
ignoring variation in RFI among individuals. Furthermore, values from feeding tables overestimate true intakes if the gas
sampling apparatus employed in the SF6 technique interferes with grazing and reduces voluntary intake. This is especially
true with young animals, where actual intakes will be less than predicted, and calculated CH4 yields will be low.
Inaccurate estimates of intake from grazing ruminants, together with uncertainties concerning CH4 measurement by SF6 ,
demonstrate a need for considerable care when using these data to estimate CH4 yield, and the validity of some published
data is questionable. An exception may be the high release SF6 capsules (170 mg/d) used by Hegarty et al. (2007), but these
have not yet been compared with calorimetry. Methane yields from individual animals based on SF6 and indirect or calculated
intakes are not defensible and should not be used to compare individuals. Methane production, or CH4 yield, of individuals
selected for either low RFI, or in response to supplementation with mitigants such as tannins, oils or saponins (Beauchemin
et al., 2008) need to be evaluated in respiration calorimeters. This technology also poses challenges because animals need
to be conned for 23 d, and feed intakes often do not reect those during normal activity. Additional problems occur when
fresh forages are offered, as cut forages do not offer the same opportunities for animal selection as occurs during grazing.

6. Systems efciency

Ruminant meat production systems have a high Ei relative to pig and poultry production (DEFRA, 2008; Table 5), and
ruminant production must strive to reduce Ei in order to expand its contribution to the global food supply in an emerging
global C economy. Ruminant farming system models have examined opportunities for limiting emissions while maintaining
constant or increased production of beef (Charmley et al., 2008; Hunter and Neithe, 2009), dairy (Lovett et al., 2006; Beukes
et al., 2010) and sheep (Alcock and Hegarty, 2006; Cruickshank et al., 2009). These papers address the impact of changing
many of the variables in Table 1 (e.g., reproduction, feed quality, potential longevity) on emissions, prot and product output.
The common message derived from assessments is that a suite of management options can increase feed efciency and
reduce the Ei of ruminant enterprises by reducing the proportion of feed energy expended on animal maintenance and
increasing the proportion expended on production. These options include increasing reproductive rate (Bentley et al., 2008),
growth rate (Hunter and Neithe, 2009) or milk production of ruminants (Beukes et al., 2010) as well as selection for RFI. While
genetic improvement is a key tool to improving these aspects of performance, superior genetics must be combined with
a level of nutrition adequate to support these high performance genetics. For example, poor nutrition will cause extended
post-partum anoestrus in cows of high genetic merit for milk, necessitating more replacement heifers and thus increasing
herd maintenance feed needs and reducing overall feed use efciency (Lovett et al., 2006).
While management changes can reduce the amount of feed required to meet a desired production target, their impact on
total CH4 emissions arising from an enterprise depends on the farmers response to having spare feed available as a result of
more feed efcient livestock. Increasing animal numbers in order to continue to use all available feed is option, but another
is to keep animal numbers unchanged, and not graze the spare pasture but harvest it or utilise the land for conserved
feed production or non-animal uses. When spare feed arising from improved feed use efciency is utilised by increased
stock numbers, emissions will increase, even though Ei will be reduced (Alcock and Hegarty, 2011). When additional feed is
provided to enable still higher production and further reduce the proportion of feed energy used in maintenance, reductions
in Ei of lambs can exceed 50%, but Ei reduction across the ock will be less than 5% due to the ewe being the major emitter
of CH4 in the production cycle (Alcock and Hegarty, 2006; Alcock et al., 2008; Cruickshank et al., 2009).
Assuming moderate rates of adoption by farmers and moderate selection pressure, Alford et al. (2006) demonstrated that
selection for improved RFI will reduce CH4 emissions from the Australian beef herd by 568 kt over 25 years. This assumes
spare feed is not utilised and animal numbers remain unchanged. It may be expected that graziers would often increase
stocking rate to utilise spare feed and, in this case, having animals with an RFI that is 10% superior than normal would only
reduce total ock emissions by <6% (Alcock and Hegarty, 2011).
Relatively simple changes in farm management, such as supplementary feeding practices, which increase productivity
of individuals will generally reduce Ei, but they are generally of less benet when calculated across the ock/herd than for
slaughter or milking animals. However, by reducing feed utilised for animal maintenance, and opting to maintain an equal
or reduced number of more productive animals, enterprises can maintain product output with reduced total emissions.
 G.C. Waghorn, R.S. Hegarty / Animal Feed Science and Technology 166167 (2011) 291301 299

If however, farm enterprises choose to increase animal numbers to use spare feed resulting from more efcient animals,
substantial increases in product output will be associated with lower Ei, but with increased total emissions. This is especially
the case if increased N fertiliser is used to achieve higher pasture DM production (Bassett-Mens et al., 2009).

7. Conclusions

Greenhouse gas emissions from ruminant animal production systems will be lowered by selecting individuals with a low
RFI because they require less feed than the average for the same level of production. Most published studies do not suggest
differences in CH4 yield for animals with divergent RFI, with lower emissions being a consequence of lower intakes. Animals
selected for low RFI can be used in both intensive and extensive farming, and should complement selection for improved
health, productivity and protability. However, individuals selected for RFI in one environment might not exhibit the trait in
another, as shown by the failure of Holstein cows selected under a total mixed ration regimen to adapt to competitive grazing
(Macdonald et al., 2008). It is essential that selection for low RFI and/or low CH4 yield, is not associated with weaknesses
in animal functional traits, such as susceptibility to disease or inferior reproduction, because animal health and fertility
are integral to efcient animal production systems. Emissions intensity is a useful measure to assess livestock production
systems from a GHG emissions perspective, especially when they are based on life cycle analysis because the real value of
ruminants as convertors of forage into high quality human foods will be realised.

Conict of interest statement

None.

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