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Perennial ryegrass breeding in New Zealand: A dairy industry perspective

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DOI: 10.1071/CP11282

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Crop & Pasture Science, 2012, 63, 107–127 Review
http://dx.doi.org/10.1071/CP11282

Perennial ryegrass breeding in New Zealand: a dairy

industry perspective

Julia M. Lee A,D, Cory Matthew B, Errol R. Thom A, and David F. Chapman C
A
DairyNZ, Private Bag 3221, Hamilton 3240, New Zealand.
B
Massey University, Private Bag 11-222, Palmerston North 4442, New Zealand.
C
DairyNZ, PO Box 160, Lincoln University, Christchurch 7647, New Zealand.
D
Corresponding author. Email: julia.lee@dairynz.co.nz

Abstract. Genetic improvement programs for livestock and pasture plants have been central to the development of the
New Zealand (NZ) pastoral industry. Although genetic improvement of livestock is easily shown to improve animal
production on-farm, the link between genetic improvement of pasture plants and animal production is less direct. For several
reasons, gains in farm output arising from improved plant performance are more difficult to confirm than those arising from
livestock improvement, which has led to some debate in the livestock industries about which plant traits to prioritise in future
breeding programs to deliver the greatest benefit. This review investigates this situation, with the aim of understanding how
genetic improvement of perennial ryegrass (Lolium perenne L.), the predominant pasture grass, may more directly contribute
towards increased productivity in the NZ dairy industry. The review focuses on the dairy industry, since it is the largest
contributor to the total value of NZ agricultural exports. Also, because rates of pasture renewal are greater in the dairy
industry compared with the sheep and beef industries, genetic gain in pasture plants is likely to have the greatest impact if the
correct plant traits are targeted. The review highlights that many aspects of ryegrass growth and ecology have been
manipulated through breeding, with evidence to show that plant performance has been altered as a result. However, it is not
clear to what extent these gains have contributed to the economic development of the NZ dairy industry. There are
opportunities for breeders and scientists to work together more closely in defining economic traits that positively influence
pasture performance and to translate this information to objectives for breeding programs, systematically linking information
on the measured traits of ryegrass cultivars to economic values for those traits to assist farmer decision-making regarding
the most appropriate cultivars to use in their farm system, and better defining genotype  environment interactions in
key productivity traits of modern ryegrass cultivars. Changes in priorities for investment of public- and industry-good funds
in forage improvement research and development will be needed if these opportunities are to be captured.

Additional keywords: breeding objectives, genetic gain, Lolium perenne, pasture performance, plant breeding.

Received 11 October 2011, accepted 15 February 2012, published online 22 March 2012

Introduction and development of perennial ryegrass
within New Zealand
Missionaries and settlers from Europe brought perennial ryegrass
(Lolium perenne L.) seed to New Zealand (NZ) in the early 19th
Century. The emergence of locally adapted ecotypes followed,
with one particularly persistent ecotype identified in the Hawkes
Bay Region on the east coast of the North Island of NZ
(Easton 1983). Since the original European germplasm tended
to be winter-dormant, an early goal of plant breeding was to
improve winter growth in NZ’s warmer climate. Pedigree lines
of the ‘Hawkes Bay’ ecotype were developed by selecting
within superior populations, ultimately resulting in a pedigree
strain with improved winter and spring herbage growth, which
was certified in 1936, and renamed Grasslands Ruanui in 1964
(Easton 1983).
Hybridisation of perennial ryegrass with Italian ryegrass
(Lolium multiflorum Lam.) was identified as another method of
improving cool-season growth, leading to the development
of the world’s first hybrid ryegrass, Grasslands Manawa,
which was released in 1943 (Corkill 1953; Easton 1983;
Rumball 1983). Grasslands Manawa was later backcrossed
with perennial ryegrass to produce a long-rotation ryegrass,
Grasslands Ariki, which had greater winter productivity, and
persistence approaching that of perennial ryegrass (Barclay
1963).
Progress was also made when it was recognised that a so-called
‘Mangere’ ecotype of perennial ryegrass from the property of
Mr Trevor Ellett (located in south Auckland in the north of the
North Island) outyielded Grasslands Ruanui and Grasslands
Ariki in on-farm experiments (Easton 1983). The perennial
ryegrass cultivars Grasslands Nui and Ellett were the first
derived from this ecotype, with certification occurring in 1975
and 1980, respectively. Many important ryegrass cultivars have
since been produced from the Mangere ecotype germplasm,

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108 Crop & Pasture Science
including Yatsyn 1, Dobson, and Bronsyn (Stewart 2006). More
recently, progress has been made with the introduction of
winter-active germplasm from north-west Spain, a region with
an isoclimatic zone similar to the North Island of NZ. Germplasm
from this region has been used in a large number of the winter-
active, late flowering ryegrasses bred in NZ, and will likely
continue to be a crucial resource in the future (Lancashire 2006).
In the early 1980s, the importance of fungal endophyte
(Neotyphodium spp.) was discovered for both animal health
and insect resistance (Fletcher and Harvey 1981; Mortimer
et al. 1982; Prestidge et al. 1982). As endophyte is seed-
borne, it has had an enormous influence on ryegrass cultivar
development, with breeders needing to ensure that appropriate
strains of endophyte are available in commercial cultivars to
provide protection from widespread insect damage in NZ. The
ecological complexity of pastures containing ryegrass, clover,
and endophyte remains a challenge for breeders attempting to
enhance animal performance by improving one component of the
system, ryegrass, as it may only explain a portion of the potential
animal productivity of the ecosystem.
Ryegrass breeding strategies
As ryegrass is cross-pollinated with a high degree of self-
incompatibility, the resulting ‘synthetic’ cultivar is very
different to self-pollinated cereal cultivars. Ryegrass cultivars
are populations of related, but genetically different, individuals
descended from several carefully chosen parent plants. The
ryegrass breeding process typically takes 10–15 years from
concept to release of a new cultivar, representing an
investment of millions of dollars.
Numerous strategies are used by ryegrass plant breeders
to develop new cultivars. The general process begins with
phenotypic screening of tens of thousands of plant genotypes
for several years, usually in the field, under cutting or grazing
and other stresses. These plants could be the first generation of
plants produced by the cross-pollination of two parent plants
differing in a particular trait (F1), F2 plants produced from inter-
pollination of F1 plants, germplasm accessions, or survivors
from old pasture. The next steps in development may include
further evaluation of selected elite plants, production of synthetic
generation 1 (Syn 1) seed based on chosen parent plants, progeny
testing, production of synthetic generation 2 (Syn 2) seed to
prevent hybrid vigour biasing experimental results, and regional
on-farm experiments with Syn 2 seed for 2–3 years under cutting
or grazing. The testing methodologies vary depending on the
purpose of the experiments and seed availability. Usually in the
early stages of the breeding process, seed supply is limited, so
experiments are restricted to replicated rows or small replicated
monoculture plots. As lines advance through the system and more
seed becomes available, evaluation is done in larger plots, with
greater replication and more sites. A substantial number of these
breeding lines that are evaluated each year never reach the market.
Current market demand that new cultivars contain an
appropriate endophyte (i.e. one that provides suitable insect
control for enhanced plant survival) presents a challenge in
breeding perennial ryegrass that adds to development timelines
and cost. Inoculating multiple new endophytes into breeding lines
adds at least 4–5 years to the breeding process, with 1–2 years
J. M. Lee et al.
required to produce sufficient seed to test, and at least a further
3 years to assess the new combinations. This situation is made
more complex because of ryegrass cultivar  endophyte strain
interactions (Easton 2007), which mean that the performance
of each ryegrass line with a particular endophyte strain is not
completely predictable and must be tested. In addition,
incorporating an effective endophyte into a breeding line is not
necessarily a simple process. For example, since the 1990s
there have been numerous papers on the potential of Festuca
endophytes, which produce lolines, an alkaloid with broad-
spectrum anti-insect effects (Dahlman et al. 1997; Popay and
Lane 2000; Popay et al. 2003; Jensen et al. 2009). Despite the
research effort, however, no ryegrass cultivar has been released
with a stable, loline-producing endophyte.
Hybridisation and introgression breeding
Hybridisation and introgression breeding are common plant-
breeding strategies, as the progeny of genetically divergent
parents often display enhanced performance or hybrid vigour,
although this effect may diminish during subsequent generations
(McClain 1982). Hybridisation involves crossing plants of two
species, normally within the same genus. In the first generation,
the inter-species hybrid produced will have an even mix of
parental genes from the two species; this balance will change
in subsequent generations depending on what the interspecific
hybrid is subsequently crossed with. In introgression breeding,
the inter-species hybrid is repeatedly backcrossed with one of the
parent species, resulting in a complex mixture of parental genes.
The genetic diversity within the outbreeding species Lolium
and Festuca provides ample opportunities for hybridisation
and introgression breeding (Humphreys 2005; Stewart 2006),
producing commercial cultivars such as Banquet II, Delish,
Grasslands Ohau, Grasslands Supreme Plus, Grasslands
Sterling, and Maverick GII (perennial ryegrass  Italian
ryegrass); and Matrix, Ultra, and Revolution (perennial
ryegrass  meadow fescue (Festuca pratensis Huds.)).
Barrett et al. (2010) created semi-hybrid perennial ryegrasses
using different cultivars and ecotypes. Annual herbage yields of
the progeny from cultivar  cultivar crosses were, on average,
1.6% greater than the better producing parent, although
there was a substantial range (+7% for Bronsyn  Revolution
to –0.4% for Bronsyn  Aries HD). Interestingly, the crosses
between cultivars with divergent heading dates demonstrated
greater hybrid vigour than those with similar heading dates. This
study indicates that semi-hybrid breeding systems warrant further
investigation.
Molecular breeding technologies
Molecular breeding technologies may provide for more rapid
and targeted cultivar development (Humphreys 2005). The
selection of superior genotypes in the field is often restricted
by the variability of environmental factors and genotype 
environment (G  E) interactions (Koornneef and Stam 2001).
This is particularly true for traits related to yield, with such traits
often controlled by numerous genes. Marker-assisted selection
(MAS) relies on identification of markers at, or closely linked to,
genes of interest that can be used to indirectly select for desirable
traits (Williams et al. 2007). A necessary precursor to MAS is to
Perennial ryegrass breeding in New Zealand
identify quantitative trait loci (QTLs) for the target traits and to
show that selection on those QTLs will shift the mean for
that trait. Numerous QTLs have been identified for a range of
ryegrass traits (e.g. Sartie 2006; Anhalt et al. 2009; Byrne et al.
2009; Khaembah 2009).
Although the tools and expertise are available to identify
beneficial QTLs, it remains to be seen how effectively these
can be applied to MAS to support plant breeders and accelerate
the development of improved cultivars (Williams et al. 2007).
One of the difficulties with outbreeding species such as Lolium is
the narrow genetic base of mapping families, so only a fraction of
the variation within the species can be described by one mapping
family, and in different mapping families, different regions of
the genome may be identified as controlling a desired trait
(Humphreys et al. 2006). This was evident when comparing
crown rust QTLs, where QTLs were identified on linkage group
(LG) 2 (Dumsday et al. 2003; Muylle et al. 2005a); LGs 1 and 2
(Muylle et al. 2005b); LGs 1, 4, and 5 (Schejbel et al. 2007); and
LGs 2, 3, 6, and 7 (Sim et al. 2007).
Molecular breeding techniques are being widely applied to the
creation of novel variation in a range of pasture plant traits related
to dry matter (DM) yield, nutritive value, and tolerance of biotic
and abiotic stresses. Examples of specific objectives have been
presented in numerous reviews (e.g. Yamada et al. 2005; Ran
et al. 2007; Humphreys et al. 2010; Kolliker et al. 2010), and
will not be discussed further here. Molecular techniques clearly
offer great promise for breaking genetic linkages between traits
in ways that cannot be achieved by other breeding strategies.
Despite the high level of investment in NZ (~$8 million),
however, genetically modified (GM) pasture plants appear to
be at least 10-20 years away from commercialisation, taking into
account the time required to ingress novel traits into well-adapted
germplasm, and to complete the necessary field evaluation and
regulatory testing procedures. A large question mark also remains
over the social and market acceptability of using GM products in
the NZ dairy industry, especially given widespread current public
concerns about the effects of intensification of dairying on NZ’s
‘clean and green’ image, and the fact that >95% of milk produced
in NZ is exported as dairy products to countries worldwide, many
of which do not currently accept imports containing, or produced
through the agency of, GM organisms.
Cultivar testing
In contrast with countries in the UK and Europe where
compulsory cultivar testing schemes are still in operation,
Table 1. Annual herbage dry matter production gains in New Zealand perennial ryegrass cultivars
Benchmark cultivar/s No. of experiments/ cultivars
Grasslands Nui 7 experiments/8 cultivars
Multiple 17 experiments/16 cultivars
Grasslands RuanuiB 8 experiments/3 cultivars
Grasslands Nui 1 experiment/7 cultivars
Average
A
In mixed swards, only the perennial ryegrass yield was used to evaluate genetic gain.
B
Results from this experiment are likely confounded by endophyte, with comparisons made between Grasslands Ruanui containing low endophyte infection,
Grasslands Nui a variable level of endophyte infection, and Ellett and Yatsyn 1 consistently high levels of standard endophyte.
Crop & Pasture Science 109
NZ’s compulsory cultivar testing scheme was abandoned in
the 1980s as part of government cost-cutting measures.
A voluntary co-operative cultivar-testing process is maintained
by the plant-breeding companies under the auspices of the NZ
Plant Breeding and Research Association (NZPBRA), called the
National Forage Variety Trials (NFVT), which determine
herbage yield over 3 years, plus ryegrass presence or absence
by point analysis at the end of the 3 years. Due to the high cost
of mixed sward experiments and their complexity, NFVT are
usually sown as pure ryegrass swards. While pure sward
experiments provide a useful measure of genetic gain in
ryegrass yield per se, pastures on-farm are usually sown with
clover (and potentially other species). Ryegrass cultivars
and different ryegrass–endophyte combinations differ in their
competitiveness and compatibility with clover (Camlin 1981;
Collins and Rhodes 1989, 1990; Sutherland and Hoglund 1989;
Sanderson and Elwinger 1999); therefore, their influence on
animal production cannot be easily estimated from pure sward
experiments, leading to the question of whether the NFVT should
include clover to improve their practical value.
Past plant breeding objectives
Herbage yield
Annual herbage yield
For several decades, a major focus of perennial ryegrass
breeding has been high annual herbage DM production (Kerr
1987), probably because harvestable DM yield was believed to be
the principal factor influencing animal production per hectare
(Williams et al. 2007). Reviews (Woodfield 1999; Easton et al.
2002) on genetic gains from plant breeding concluded that
perennial ryegrass herbage production has increased by ~0.5%
per year (Table 1). These results are similar to those from France
and Italy (0.25%; Allerit 1986; Veronesi 1991), Europe and the
Netherlands (0.3%; Van Wijk and Reheul 1991; Sampoux et al.
2010, 2011), Belgium (0.3–0.5%; Van Wijk and Reheul 1991;
Chaves et al. 2009), and the UK (0.6%; Aldrich 1987). Depending
on the country and report, the genetic gain in perennial ryegrass is
also similar to, or about half, that of other pasture species such as
white clover (Trifolium repens L.: 0.6%, Woodfield and Caradus
1994; 1.2–1.5%, Woodfield 1999), Italian ryegrass (1.2%, Easton
et al. 1997; 0.2–0.5%, Brummer et al. 2009), and tall fescue
(Festuca arundinacea Schreb.: 0.1–0.6%, Brummer et al. 2009).
Data from the NFVT (Fig. 1) indicate that herbage DM
production gains are still being achieved with new ryegrass
Pure/mixed swardsA Genetic gain Reference
(%/year)
Pure/mixed 0.25 Pennell et al. (1990)
Pure/mixed 0.40 Easton et al. (2001)
Mixed 0.60 Kerr (1987)
Mixed 0.73 Thom et al. (1998)
0.50
110 Crop & Pasture Science
0 2000 4000 6000 8000 10 000 12 000 14 000 16 000
Ryegrass yield; kilograms of dry matter per hectare
Fig. 1. Annual yields of diploid ryegrass cultivars in New Zealand under
high nitrogen supply (source: NZPBRA 2010). Cultivars whose error bars do
not overlap are significantly different at P = 0.05. Each cultivar was involved
in at least three completed approved experiments, with at least one of
those experiments conducted north of Taupo in the North Island of New
Zealand. Experiments lasted for 3 years, plus the establishment autumn.
Unless specified, all cultivars contained standard endophyte. Supreme Plus,
Revolution, and Matrix are hybrid ryegrasses; the rest are perennial.
Uncertified LP is uncertified perennial ryegrass.
cultivars. Averaged across multiple experiments, the top-yielding
cultivar was Ceres One50, which had a 2007 Plant Variety
Rights application date. Ceres One50 yielded ~10% more than
Grasslands Nui, a 1970s cultivar which is often used as a
benchmark for calculating genetic gain. It is clear, therefore,
that breeders are continuing to make genetic progress in ryegrass
yields when evaluated as a pure sward under high nutrient supply.
The rate of genetic gain in perennial ryegrass annual DM
yield is often compared with that in cereal or grain crops, and
criticised for being lower. However, this is not surprising
considering that (1) the changes in harvest index are limited,
i.e. all aboveground mass is harvested compared with crops in
which the proportion of grain can be manipulated; (2) ryegrass
yield is measured following repetitive defoliations/grazings over
successive years, rather than as seed at maturity in annual crops;
(3) compared with other crops, a greater range of beneficial traits
have been selected for ryegrass, rather than just herbage yield
alone; and (4) ryegrass is a perennial with a longer breeding cycle
(Camlin 1997; Humphreys 1997; Woodfield and Easton 2004;
Barrett et al. 2010; Stewart and Hayes 2010).
Although annual herbage DM yield of a forage grass is
important, it is only a partial indicator of merit, as the ultimate
J. M. Lee et al.
value of perennial ryegrass lies in the resulting animal production
from the mixed sward (Munro et al. 1992; Laidlaw and Reed
1993; Reed 1994). There is a paucity of animal production data
to validate gains in herbage production, mainly due to the
experimental costs involved. Experiments initiated by the late
Dr R. W. Brougham indicated that weaner bulls grazing new
perennial ryegrass, hybrid ryegrass, and clover pastures had
greater liveweight gain than those grazing old permanent
pasture (1965 v. 1761 kg liveweight/ha, respectively; Cosgrove
et al. 2003). Unlike the new pastures, however, the old pastures
were not cultivated and resown, making it difficult to separate
the benefit of genetic gain through plant breeding from the
beneficial effects of cultivation (e.g. weed and pest control,
enhanced mineralisation of soil nitrogen). This highlights just
one of the difficulties with cultivar evaluation, the necessity of
including a valid control. The experiment does, however, clearly
indicate the benefits of pasture renewal to farmers.
Crush et al. (2006) compared mixtures of perennial ryegrass
cultivars available in the 1980s (Ellett, Grasslands Nui, and
Yatsyn 1) with those available in the late 1990s (Aries HD,
Bronsyn, and Grasslands Samson). Overall, no effect of cultivar
mixture on annual herbage production was measured, with
yields averaging 17.2  0.9 t DM/ha.year over three full years,
nor was there any significant impact of ryegrass cultivar on
milk or milk solids yield per cow, milk solids yield per
hectare, or the economic farm surplus. One interesting result
was that the tiller density of the 1980s ryegrass pastures declined,
likely due to the lower endophyte infection allowing greater
insect damage (Eerens et al. 2001). This reduction in tiller
density allowed a greater proportion of white clover in the
swards containing 1980s ryegrass/1990s clover compared with
1990s ryegrass/1990s clover (32 v. 22%, respectively; Eerens
et al. 2001), probably contributing to the increased milk
production during the first year (Woodward et al. 2001), and
highlighting the complex interactions between grass, endophyte,
and clover in mixed swards.
The results from the experiment of Crush et al. (2006) are
often debated, questioning whether plant breeding has failed to
improve pasture yield, whether the experimental design lacked
sufficient sensitivity to identify real differences between cultivar
mixtures, or whether different levels of endophyte infection
confounded the results. Data from the NFVT indicate that
ryegrass yield gains in monocultures under high nitrogen (N)
supply are real (e.g. Fig. 1), but the gains expected from 1990s v.
1980s cultivars based on best estimates from NFVT are less than
the least significant difference for the experiment. In addition,
G  E interactions and interactions between ryegrass and other
plant species may have meant that cultivar differences measured
in monocultures under high N supply were lost in mixed pastures
under lower N supply. Regardless of the interpretation, this study
provides valuable lessons for future cultivar evaluation. First,
experimental treatments must be sufficiently divergent so that
any differences, or lack thereof, will be indisputable. In an
animal strain experiment, cows with 1970s genetics were bred
specifically for comparison with 1990s genetics. This provided
suitably divergent groups, which allowed the clear gains in milk
and milk solids yields to be detected (Macdonald et al. 2008).
Second, cultivars should be tested in a range of environments
to elucidate G  E interactions. Third, endophyte infection
Perennial ryegrass breeding in New Zealand Crop & Pasture Science 111

levels and strains ideally should be similar; this is likely to be
difficult to achieve in future evaluations, as current endophyte
strains are different to those of 25 years ago.
Seasonal distribution of herbage yield
and heading date
While increased annual herbage production is important, more
closely aligning herbage production and animal requirements
during key periods (e.g. late winter/early spring and summer/
autumn) is also valuable (Woodfield 1999; Easton et al. 2002).
This was confirmed by Brookes et al. (1993), who demonstrated
that while annual DM yields of three diploid ryegrass cultivars
(Grasslands Nui, Yatsyn 1, and Embassy) were similar, Embassy
produced 10% more feed between January and August, and 5%
less feed between September and December. Computer models
estimated that this pattern of growth would allow a stocking rate
4–6% greater than the other two cultivars, resulting in greater
milkfat production, reduced requirements for supplementary
feed, and a 5–7% increase in gross margin. Unfortunately,
however, on-farm experiments have not been undertaken to
confirm these results.
Gains in herbage yield during key seasons have been
demonstrated, with late 1980s/early 1990s cultivars (Vedette,
Dobson, Embassy, and Grasslands Impact) producing 10–15%
more during summer than Yatsyn 1, which was bred in the
early 1980s (Thom et al. 1998). More recently, increases in
summer–autumn yields of up to ~1 t DM/ha have been
reported, depending on the benchmark and new cultivars,
endophyte, and the testing regions (Easton et al. 2002;
NZPBRA 2010).
One of the methods by which seasonal distribution of yield
may be manipulated is by altering the heading date. Midseason-
maturity NZ perennial ryegrass cultivars such as Grasslands
Ruanui, Grasslands Nui, Ellett, and Yatsyn 1 have similar
heading dates, with emerged seedhead on 50% of plants by
around 22 October each year (Day 0, Fig. 2). Over time, plant
breeding has produced cultivars with a range of heading
dates from 17 days before day 0 (cv. Meridian) to 25 days
after day 0 (cvv. Bealey, Grasslands Halo, Grasslands Sterling,

Early heading DAY

date –17 Meridian

–6 Embassy, Vedette

–3 Banks, Ceres Kingston

Mid-season –1 Ceres Cannon

heading date 0 Bronsyn, Ellett, G. Kamo, G. Nui, Marathon, Yatsyn 1
1 G. Commando, G. Pacific, G. Ruanui
2 Aries HD
3 Extreme, G. Samson
4 G. Hillary

7 Arrow, Dobson
8 G. Ohau (t)
9 Delish (t)

12 Marsden, Perun (t)

13 Harper
Late heading 14 Alto
date 15 AberDart, G. Supreme Plus
16 Trojan
17 Maverick GII
18 Banquet II (t)
19 AberMagic, Revolution
20 Ceres One50
21 Expo, G. Impact, Ultra
22 Base (t)
23 Matrix
Very late
25 Bealey (t), G. Halo (t), G. Sterling (t), Quartet II (t)
heading date

Fig. 2. Heading dates of New Zealand diploid and tetraploid (t) perennial, long rotation, and
hybrid ryegrass cultivars. Day 0 is typically 22 October, but this can vary between years and under
different grazing management. G., Grasslands.
112 Crop & Pasture Science J. M. Lee et al.

and Quartet II). Earlier heading cultivars tend to have greater DM flowering aspects, developing late-heading ryegrasses that
growth during late winter/early spring, whereas later heading maintain their ‘leafiness’ longer and retain better nutritive
cultivars retain better herbage nutritive value during late spring value during late spring (Easton et al. 2002), as well as
(Easton et al. 2002). cultivars which have reduced aftermath heading (Forde et al.
The effect of heading date on animal production has been 1988). Evaluating the impact of altered flowering behaviour on
investigated in Ireland (Gowen et al. 2003; O’Donovan and nutritive value is, however, difficult as there is a lack of primary
Delaby 2005); it is assumed that these findings may be data, particularly comparing perennial ryegrass cultivars.
relevant to NZ due to similarities between the two pasture-
based systems. O’Donovan and Delaby (2005) demonstrated Leaf strength
that there was no effect of heading date on milk production at
A divergent selection was carried out on Grasslands Ariki to
high stocking rate. At low stocking rate, however, milk
form selections of high and low leaf tensile strength, with
production from cows grazing late-heading ryegrasses (first
the hope that lower leaf tensile strength and associated lower
half of June) was 5% greater than those grazing cultivars with
cellulose content would improve animal performance (Wilson
intermediate heading dates (last half of May, i.e. classified as
1965). This eventually led to sheep production experiments
late-heading in NZ). In addition to the increased milk production
which, unfortunately, were confounded by endophyte effects
from late-heading ryegrasses, milk protein, lactose (Gowen et al.
(Lancashire and Ulyatt 1975; Lancashire et al. 1977).
2003; O’Donovan and Delaby 2005), and fat (Gowen et al. 2003)
However, the selection with low leaf tensile strength was
yields were also greater. Factors contributing to the increased
released as ‘Grasslands Marsden’ in 1989.
production from cows grazing ryegrasses with late heading dates
A further selection for leaf shear strength was made from
included greater pre-grazing herbage mass, herbage allowance,
a population of Grasslands Nui plants and L. perenne 
and herbage intake, and increased digestibility (O’Donovan and
(L. perenne  L. multiflorum) hybrids (Easton 1989). The
Delaby 2005). As there is less supplementary feeding in NZ than
plants with low leaf shear strength had shorter, narrower
in Ireland, as well as a constant, year-round stocking rate, an
leaves with 27% less resistance to shear per unit cross-section
investigation of how heading date affects animal production in
area than the plants with high leaf shear strength (Inoué et al.
NZ would be valuable. Potential value could be demonstrated
1994a). Sheep grazing the selection with low leaf shear strength
using growth rate data in a computer model, followed by on-farm
had increased rates of DM ingestion, but DM intakes (DMI)
validation experiments if differences were indicated.
and liveweight gain were not consistently increased (Mackinnon
et al. 1988; Inoué et al. 1994b), leading to the conclusion that
Herbage nutritive value ‘selection for reduced leaf shear breaking load per se did not
Herbage nutritive value is a complex trait, as pastures generally improve feeding value’ and that associated changes in leaf
consist of perennial ryegrass, white clover, and potentially other morphology need to be taken into account.
plant species. Therefore, improving the nutritive value of ryegrass
alone may not affect the actual nutritive value of the pasture, or Tetraploids
animal production, as much as expected. Nutritive value of the
Perennial ryegrass plants exist naturally as diploids, with each
pasture as a whole is influenced strongly by the proportion of
cell containing seven pairs of chromosomes (2n = 2x = 14;
clover in the sward, with strong and competitive ryegrass growth
Humphreys et al. 2010). One of the earliest achievements of
(e.g. high tiller density) potentially decreasing white clover
plant breeders was to artificially double the chromosome number
growth and reducing pasture nutritive value and subsequent
to form tetraploids (Easton 1983), which have larger cells, greater
animal production (Rhodes et al. 1987; Woodward et al. 2001).
water content, and larger leaves, tillers, and seeds (Charlton and
Nutritive value traits of ryegrass have historically received
Stewart 1999). Despite the significance of this, there is little
less attention than herbage yield (Smith et al. 1997), which may
NZ research documenting the subsequent effects on pasture or
be attributable to a lack of clarity over which characteristics
animal production. In the UK, annual herbage production of six
were more desirable and/or a failure of early experiments to
tetraploid ryegrasses was greater than that from six diploids,
demonstrate significant improvements in animal production
averaging 10.9 v. 9.9 t DM/ha.year, respectively (Gilliland et al.
(Easton et al. 2002), possibly as earlier experiments were
2002). Tetraploids also had a greater WSC yield, longer extended
likely confounded by endophyte interactions. Specific ways in
tiller height, and greater bulk density and green leaf mass above
which breeders have focussed on improving herbage nutritive
the defoliation height (Gilliland et al. 2002), all factors which
value of ryegrass include altering flowering behaviour, reducing
can improve animal intake and production (Poppi et al. 1987;
leaf strength, increasing ploidy, and increasing herbage water-
Hodgson and Brookes 2000; Parga et al. 2000; Miller et al.
soluble carbohydrate (WSC) content; the impact of these will be
2001b).
described below.
In Ireland, milk production from cows grazing two tetraploid
ryegrasses was similar to production from cows grazing two
Flowering behaviour diploid ryegrasses (Gowen et al. 2003; O’Donovan and Delaby
Nutritive value in ryegrass, and indeed any grass, is dictated 2005). In these experiments, cows grazing the tetraploids were
largely by the flowering behaviour, declining during seedhead offered 6–8% less feed due to lower pre-grazing masses, which
production. The main flowering period occurs in spring (Fig. 2); may have limited their milk production. Care should also be taken
however, there is often a second flowering period or aftermath when interpreting these results, as one of the tetraploids in
heading during summer. Breeders have focussed on both of these both experiments, Napoleon, was demonstrated to be inferior
Perennial ryegrass breeding in New Zealand
(production- and persistence-wise) to other tetraploids, and was
the only cultivar of the four to be removed from the 2006
Recommended Variety List (DAF 2006), likely reducing the
potential of the tetraploids in these experiments.
In the Netherlands, high allowances of diploid or tetraploid
perennial ryegrass herbage (35–45 kg DM/cow.day above 45 mm
stubble height) were offered to avoid confounding effects on
intake (Hageman et al. 1993). Cows offered tetraploids had a
tendency for greater intake (+0.6 kg organic matter/cow.day) and
4–5% greater milk solids production. This was likely due to a
combination of factors, including more rapid rates of organic
matter degradation, greater soluble fractions/reduced insoluble
fractions, greater WSC content, and reduced cell wall content in
tetraploids.
Thus, overseas research indicates that tetraploids have the
potential to increase animal production. In addition, pastures
based on tetraploid ryegrass tend to have greater clover
content than diploid-based pastures, which is due, at least in
part, to their more erect, less densely tillered nature (Frame
and Boyd 1986). Increased clover content also enhances the
nutritive value of pastures, which may result in increased
milk production (Harris et al. 1998). One possible drawback to
tetraploids, however, is that, anecdotally, they may be less
persistent than diploids, possibly due to the fact that cows
may graze tetraploids to a lower residual height than diploids
(Gowen et al. 2003; O’Donovan and Delaby 2005) and/or that
that tetraploids may be more susceptible to insect attack (Popay
et al. 1995). Persistence may be less of a problem in recently
released tetraploids that have tiller numbers similar to diploids,
compared with older tetraploids with lower tiller numbers;
however, this requires further investigation. Research into
tetraploids in a NZ dairy farming system should be undertaken
to investigate their value.
Water-soluble carbohydrates
The breeding effort into high sugar grasses, i.e. grasses that
contain elevated concentrations of WSC, was initiated in the UK
based on predictions of increased milk production through
increased digestible DMI (Miller et al. 2001b). In addition, it
was predicted that grasses with increased WSC would mitigate
adverse environmental effects by improving N-capture efficiency
in the rumen, thereby increasing N supply for milk production
and reducing the amount of urinary N deposited on the soil
(Miller et al. 2001b).
Edwards et al. (2007a, 2007b) reviewed supporting evidence
for the hypothesised benefits (Miller et al. 2001b), as well as
experiments where data fail to consistently support either the
milk production responses (Miller et al. 2000, 2001a; Tas et al.
2005, 2006a, 2006b; Taweel et al. 2005; Moorby et al. 2006;
Cosgrove et al. 2007a) or increased N-use efficiency (Tas et al.
2006a). While conclusive proof of the animal production benefit
from grasses with increased WSC remains elusive, when data
from all experiments were combined, it became evident that
as the WSC : crude protein (CP) ratio increases above 0.7,
the proportion of N intake excreted in urine declines (Fig. 3;
Edwards et al. 2007a; Parsons et al. 2011).
In NZ, monthly average CP concentrations in the herbage tend
to range between 18 and 32% DM, with peaks during autumn and
Crop & Pasture Science 113
60
50
40
NUE urine
30
20
10
0
0 0.5 1.0 1.5 2.0 2.5
WSC:CP ratio
Fig. 3. Nitrogen use efficiency (NUE; g nitrogen excreted in urine/100 g
nitrogen eaten) plotted in relation to the water-soluble carbohydrate
(WSC) : crude protein (CP) ratio of the diet (source: Parsons et al. 2011).
*, 2001 data from Tas et al. (2005, 2006b); *, 2000 data from Tas et al.
(2005, 2006b); ~, Miller et al. (2000); ¤, Miller et al. (2001a, 2001b);
&, Moorby et al. (2006).
late winter/early spring (Moller et al. 1996; Roche et al. 2009).
This indicates that to achieve a desirable WSC : CP ratio of
1.5 (Parsons et al. 2011), grasses need to achieve WSC
concentrations of 27 and 48% DM, respectively. These values
are well in excess of those measured in the majority of NZ
experiments (Moller et al. 1996; Cosgrove et al. 2007a,
2007b, 2009; Hume et al. 2010). While Easton et al. (2009b)
demonstrated that ratios 1.5 were achievable in Canterbury
under moderate N fertilisation (160 kg N/ha) and longer grazing
intervals (4 weeks during peak growth, longer during the
remainder of the year), it appears unlikely that a ratio of 1.5
would be achievable by the 3-leaf stage of regrowth, the point
at which many NZ pastures are grazed, or in pastures receiving
a large amount of N fertiliser. Nevertheless, Fig. 3 indicates a
progressive decline in the proportion of N intake excreted as urine
at a WSC : CP ratio >0.7, so any further increase in this ratio, by
altering the WSC or CP content, may be valuable to the NZ dairy
industry as a whole in terms of environmental sustainability.
Seed yield
Despite the negative impact of flowering on herbage nutritive
value, seed-yielding capacity is vital in grass breeding (Elgersma
1990; Cunningham et al. 1994) as it allows the economic delivery
of seed to farmers. Some NZ cultivars, for example ‘Tolosa’, were
unable to be successfully delivered to farmers due to low seed
production (Stewart and Hayes 2010).
Disease and pest resistance
Crown and stem rust
The most important perennial ryegrass diseases in NZ
are crown and stem rust (Puccinia coronata and P. graminis,
114 Crop & Pasture Science
respectively), which reduce ryegrass herbage and seed yield,
herbage nutritive value, root mass, plant persistence, and,
potentially, animal intake (Corkill 1956; Hunt and Easton
1989; Kimbeng 1999). Although white clover is not a host of
crown rust, research indicates that white clover yields were
also reduced in a mixture with rust-infected ryegrass (Mattner
1998; Mattner and Parberry 2001). Breeding programs have
been screening for rust resistance since the 1930s (Easton
1983), but unfortunately there are few published experiments,
making it difficult to truly demonstrate the benefit of this work.
In the late 1980s, 11 ryegrass cultivars were assessed for crown
rust infection in six locations around NZ (Easton et al. 1989).
Yatsyn 1 was less affected than earlier cultivars (Grasslands
Ruanui, Grasslands Nui, and Ellett), suggesting that breeding
had been successful in improving crown rust resistance, at least
in certain cultivars. More-recent evidence confirms that while
there is some variation in crown rust resistance between different
cultivars produced after 1985, all have improved rust resistance
compared with Grasslands Nui and Yatsyn 1 (Thom et al. 1998;
Easton et al. 2002), despite the rust fungi mutating over time
and/or changes in the frequency of different rust races.
Endophyte
Research over the past 30 years has clarified many of the
issues around endophyte in perennial ryegrass (Easton et al.
2007). The deterrent effects of endophytes on insect feeding
have been identified as having a major impact on the agronomic
performance (yield and tiller survival) of ryegrass cultivars
(Prestidge et al. 1982; Popay et al. 1999; Hume et al. 2007).
Animal production may also be affected by the presence of the
standard endophyte that produces the alkaloids ergovaline and
lolitrem B (Gallagher et al. 1981; Prestidge 1993). Ergovaline in
ryegrass has been linked to heat stress in sheep (Fletcher et al.
1999), although no direct association has been demonstrated with
cattle (Bluett et al. 2005), while lolitrem B causes ryegrass
staggers and can reduce milk production (Fletcher and Harvey
1981; Thom et al. 1999, 2010).
Substantial plant breeding effort over the last 15–20 years
has gone into discovery, selection, and inoculation of novel
endophytes into ryegrass with reduced toxicity towards
animals, while trying to maintain insect deterrent capability
similar, or superior, to standard endophyte (Easton et al.
Table 2. Ryegrass cultivar ¾ endophyte combinations available for farmers
(t), Tetraploid; G., Grasslands
Endophyte Alkaloids produced
Standard Peramine, lolitrem B, ergovaline
NEA2A Peramine, lolitrem B, ergovaline
Endo5 (also Peramine, ergovaline
called AR5)
AR1 Peramine
AR37 Epoxy-janthitrems
A
NEA2 is a mixture of two distinct genotypes or variants (van Zijll de Jong et al. 2008). It is possible that NEA2-infected cultivars may contain different proportions
of the two genotypes, which may in turn affect performance of the cultivar; this requires further investigation.
J. M. Lee et al.
2007). This process, carried out by plant breeding companies
and AgResearch, has resulted in new ryegrass endophytes being
available to farmers (NEA2, Endo5, AR1, AR37; Table 2).
Agronomic testing of the new selections continues to provide
evidence of the success or otherwise of the selected endophytes in
particular cultivars and in different regions (Bluett et al. 2003,
2005; Hume et al. 2009; Popay and Thom 2009; Thom 2010).
Different endophytes provide differing levels of protection
against certain insect species, and this can have a substantial
impact on summer survival under stressful conditions (Thom
2010). Of the endophytes currently available to farmers, AR37
provides good control against many insect pests (e.g. black beetle
adults, pasture mealy bug, root aphid, porina, and Argentine stem
weevil larvae), and resulted in pastures with the greatest tiller
density after the summer 2008 drought in the Waikato. Of the
other novel endophytes available, Endo5 provides good control
against black beetle adults, pasture mealy bug, and Argentine
stem weevil larvae; AR1 controls attack from Argentine stem
weevil and pasture mealy bug; and NEA2 provides good control
of black beetle adults and moderate control of Argentine stem
weevil larvae in the tetraploid cultivar Bealey (Popay and
Hume 2011). It is important, therefore, for farmers to select an
appropriate cultivar with an endophyte that will control local
insect populations (e.g. use AR37 where black beetle and root
aphids are a problem; Popay and Thom 2009) and that the seed
received by the farmer contains a high viable endophyte infection
level (>70%).
Persistence
Persistence is often defined as the survival of a desired species
through time without major intervention (e.g. Clark 2011). It is
a particularly complex plant trait, with numerous contributing
factors. Clark (2011) categorised these factors into a primary
subset leading to a progressive decline in plant density, and a
secondary subset that will rarely start a decline in persistence
alone, but can lead to catastrophic plant losses in weakened
pastures on interaction with other primary and secondary
factors. The primary factors included soil type (water-holding
capacity and texture), summer rainfall, plant nutrient status,
and plant population survival mechanisms (tiller dormancy,
birth and death rates, seed production and establishment,
endophyte status), while secondary factors included pests,
Cultivars available
Ceres Kingston, Grasslands Pacific, Matrix
Bealey (t), Trojan
Banquet II (t), Quartet II (t)
AberDart, AberMagic, Alto, Arrow, Bronsyn, Ceres One50, Delish (t), Expo, Extreme,
G. Commando, G. Hillary, G. Impact, G. Ohau (t), G. Samson, G. Sterling (t), G. Supreme Plus,
Harper, Helix, Meridian, Revolution, Revolution Ultra
Alto, Base (t), Ceres One50, Extreme, G. Commando, G. Halo (t), G. Kamo, G. Ohau (t),
G. Samson
Perennial ryegrass breeding in New Zealand
diseases, weeds, and high grazing intensity resulting in
overgrazing, pugging, and/or pasture pulling.
Anecdotal reports suggest that old ryegrass cultivars persist
better than modern cultivars. This is not substantiated by research
(Easton et al. 2001; Crush et al. 2006; Easton et al. 2011),
although these experiments did not last more than 4 years. In
fact, L’Huillier and Aislabie (1988) state that:‘In ryegrass/white
clover dairy pastures reliance on vegetative propagation appears
to be inadequate to maintain sward stability’, indicating that
persistence of ryegrass in dairy pastures has been an issue for
more than two decades. There are several possible reasons for
the perception that newer cultivars are less persistent. First, since
the late 1970s there has been a general trend for reduced
summer–autumn rainfall in NZ (Ummenhofer et al. 2009).
This, combined with associated factors such as overgrazing
due to poor pasture growth, has likely reduced persistence of
ryegrass cultivars compared with those grown in the 1970s.
Widespread use of cultivars infected with black beetle-
susceptible AR1 endophyte in black beetle-prone areas in the
last few years may have also exacerbated the problem. The
perception may also be partly due to less natural reseeding of
modern cultivars. Most modern ryegrass cultivars bred in NZ
have reduced aftermath heading (A. V. Stewart, pers. comm.),
which may reduce seed drop and seedling recruitment. In
addition, current grazing management strategies tend to focus
on improved nutritive value by removing seedheads when they
are immature (Hoogendoorn et al. 1992). In some situations,
the lack of seedling recruitment through natural reseeding
may limit persistence, as plants are reliant either on survival of
vegetative tillers (L’Huillier and Aislabie 1988; Waller and
Sale 2001), which have a poor survival rate during dry
summers (Silsbury 1964; Waller et al. 1999), or on survival of
daughter tillers from flowering tillers (Matthew et al. 1993),
which can also be problematic.
Although ‘persistence’ is often stated as an objective of
plant-breeding programs, many claims of ‘persistence’ are
based on <5 years of data, which is arguably an insufficient
length of time to accurately evaluate persistence. Woodfield
and Easton (2004) suggest that persistence may be inversely
correlated with yield; therefore, plant-breeding programs also
run the risk of discarding potentially valuable germplasm
which may have marginally reduced annual herbage yields but
better long-term persistence. The development of evaluation
systems which substantiate persistence over a longer period is
necessary. It has been proposed that, in addition to the NFVT
DM yield protocols, ‘persistence’ experiments should be
conducted in which pre-commercial cultivars and commercial
controls are sown at sites with known persistence challenges,
and plant population survival tracked over time, as happens
already in some breeding programs. This will allow the claims
of better persistence to be proven conclusively (or not), and
should improve farmers’ perceptions and uptake of pasture
renewal.
Plant breeding in the future
With the process of cultivar development taking up to 15 years
to produce a commercial cultivar, it is important to revisit
objectives of plant-breeding programs to ensure that they
Crop & Pasture Science 115
are still relevant. Indeed, a conference involving stakeholders,
breeders, evaluators, and grassland scientists was held in Ireland
in October 2010 for this very purpose (O’Donovan and Hennessy
2010). The opening paper stated that plant breeding should focus
on seasonal growth, nutritive value (particularly during summer),
providing sward structures suitable for grazing (i.e. high
proportion of green leaf), and persistence (O’Donovan et al.
2010). These traits are probably similar to those desired within
NZ, although it must be recognised that the Irish system is less
complex, with pure ryegrass swards lacking endophyte and
clover.
Parsons et al. (2011) suggest that plant-breeding objectives
should be more focussed, translating objectives (e.g. increased
metabolisable energy) into traits (e.g. alter plant metabolism to
increase fructan content), and quantifying the target (e.g. the
required level of expression of this trait is a WSC : CP ratio of 1.5).
They cite the work conducted on low respiration ryegrass as an
example of trait selection that was initially successful (Wilson and
Jones 1982), but advantages diminished over time in subsequent
field experiments (Robson et al. 1983, 1988), due perhaps to
either the trait being lost or its expression being disguised by
G  E interactions, and so no commercial cultivar was ever
released. Parsons et al. (2011) propose that abandoned traits
such as this should be revisited with new techniques such as
gene expression profiling, to investigate the G  E interactions
that potentially limit expression of the trait and to ensure such
traits are ‘locked in’ to the plant. The remainder of this review
takes a step in this direction, identifying candidate plant traits for
such research and suggesting an integrated breeding, agronomy,
and farming systems approach for the future.
In proposing traits that could be beneficial for dairy pasture
performance in the future, we recognise that investment in
the basic research required to define the traits precisely is
beyond the resources and remit of commercial breeding
companies. Rather, it falls more in the domain of public- and
industry-funding agencies. We also note that public investment
in the underpinning plant (excluding molecular genetics), soil,
and entomological sciences in NZ has declined steadily over
the past two decades. A recent review of the forage improvement
system in NZ recommended that capability in the sciences
underpinning plant improvement should be boosted and
linked with increased effort to measure impacts upon on-farm
performance (J. Morrison, pers. comm.). The latter point is
critical because without good evidence for the impact that trait
manipulation could have on the productivity or environmental
outcomes of dairying systems, there will be little incentive for
plant breeders to shift their selection focus. To achieve this,
improved knowledge of the biophysical mechanisms
associated with specific traits must be translated into realistic
objectives for plant-breeding programs. This implies that a
closer working relationship between scientists and breeders is
essential; there is, perhaps, a sound argument for some public-
and industry-good funds to be dedicated to this purpose. Finally,
we note that there are likely to be trade-offs between diverting
resources to address ‘new’ traits, and the rate of gain that can be
sustained in other traits such as DM yield or pest and disease
resistance. Industries such as dairying may need to accept some
slow-down in progress in the short term to realise longer term
benefits.
116 Crop & Pasture Science
Persistence
Parsons et al. (2011) suggested that the failure of a new pasture to
‘persist’ as expected may be a result of:
1. Loss of plants from the population that established from the
sown seed (i.e. high rates of plant mortality, which may reflect
poor adaptive ability of the sown cultivar and/or significant
episodic lethal stress events such as drought and pest attack);
2. Loss of the specific trait contained in the cultivar that was
sown (i.e. sown plants survive, but the trait is not expressed or
is lost altogether); and
3. Loss of overall yielding ability in the pasture (i.e. sown plants
and specific traits persist, but other yield-related traits are not
expressed or are lost altogether).
Although there is considerable information regarding the first
of these factors (plant mortality) in NZ pastures, especially in
relation to ryegrass–endophyte associations and their benefits for
plant survival in the face of pest challenges (see, for example,
Popay and Hume 2011; Rennie et al. 2011), the other two factors
have received virtually no attention in agronomic investigations
into the problem of persistence. There is an urgent need for
systematic, scientific information on changes in population
biology, the genetic structure of sown populations, and plant
phenotype for different functional groups of perennial ryegrass
(e.g. diploids and tetraploids) in a range of dairying environments
in NZ. Without this information, it will be difficult to define
exactly what is not persisting when the performance of a
perennial ryegrass pasture declines after sowing. In turn, it will
not be possible to accurately identify plant-breeding objectives,
desirable traits, and suitable germplasm to address the problem,
nor to develop suitable evaluation methods to assess progress in
improving persistence. Systematic investigation of the problem
of perennial ryegrass persistence, using the framework described
above and by Parsons et al. (2011), should ensure that:
1. Robust control treatments are included, i.e. ‘older’ as well as
modern ryegrass cultivars, and perhaps other species known
to have greater persistence than perennial ryegrass in specific
environments, as discussed by Chapman et al. (2011a).
2. Critical shoot and root characteristics, such as those described
below, are measured so that the mechanisms associated with
poor or good persistence can be determined and included in
future plant screening and evaluation programs.
3. Interactions with clover are determined; for example, there
is evidence that while more-persistent ryegrasses may yield
more and reduce weed ingress, they may also be less
compatible with white clover (Camlin 1981). This should
be tested with current and new NZ cultivars and the impact on
milk production determined.
Shoot morphology
Before the 1990s, plant growth processes such as root
formation (Garwood 1967; Caradus and Evans 1977), tiller
population birth and death (Langer 1958, 1963; Jewiss 1966;
Korte 1986), and leaf formation and death (Bircham and Hodgson
1983) were often studied in isolation. Inter-relationships between
component processes of the grass shoot and ‘productive fitness’
as reflected by measures such as leaf area index were subsequently
J. M. Lee et al.
mapped (Chapman and Lemaire 1993; Lemaire and Chapman
1996), and extended as depicted in Fig. 4 (Bahmani 1999).
Bahmani’s study used the diploid cultivars Grasslands Ruanui,
a fine-leaved, densely tillered, persistent plant, and Ellett, a more
erect plant with larger leaves and tillers. Ellett had 13% greater
annual herbage production over 2 years under rotational dairy
cow grazing, with increases in growth particularly evident during
spring–summer (Bahmani et al. 2001). Factors contributing to
the superior growth of Ellett included more rapid leaf elongation
rates (LER) contributing to longer leaves and greater tiller
weight (Bahmani et al. 2000, 2001, 2002) and production of
more reproductive tillers in response to N (Bahmani et al. 2002).
Ellett also had lower tiller density/tiller numbers per plant as a
result of reduced site filling (i.e. ratio of new tillers to new leaves;
Bahmani et al. 2000, 2001, 2002, 2003) and a greater tendency
towards pulling (Bahmani et al. 2001).
Although increased yield is a desirable trait, certain
morphological traits of Ellett, such as the tillering habit (i.e.
less-densely tillered, greater production of reproductive tillers;
Bahmani 1999), were possibly less desirable than those displayed
by Grasslands Ruanui. There are conflicting views about the
effect of increased reproductive tillering on persistence. While
Bahmani (1999) believed that increased reproductive tillering
was likely to reduce persistence, Matthew et al. (1993) regarded
tillering from reproductive tillers as an alternative, rather
than inferior, persistence strategy, and Waller et al. (1999)
demonstrated that daughter tillers growing on old reproductive
stems in a summer dry environment had a greater chance of
survival than those growing on vegetative tillers (56% v. 12%,
respectively). This may have been due to greater carbohydrate
and protein content remaining in the reproductive stems
providing nourishment for the tillers during stress. These data
indicate that reproductive tillering may not always compromise
persistence; however, further investigation with current NZ
cultivars is required to fully understand this relationship.
The multiple trait interactions elucidated by Bahmani (1999)
in Fig. 4 were further explored using principal component
analysis to identify plant genotypes with combinations of traits
conferring higher productivity (Sartie 2006; Sartie et al. 2011).
The study found that tiller size was a ‘yield-neutral’ trait, not
correlated positively or negatively with plant dry weight. This
confirmed the understanding that tiller size/density compensation
provides a range of plant morphology configurations to deliver
the same leaf area, and in the field is a mechanism by which grass
swards can adapt to variation in grazing height (e.g. Davies 1988;
Matthew et al. 1995). Earlier findings of an association between
higher LER and increased plant yield (Van Loo 1993; Skinner
and Nelson 1994; Bahmani 1999) were supported by these data,
but there was also a conundrum for breeders intending to select
on LER, as there was not just a simple univariate correlation
between LER and plant yield. Rather, the relationship was only
revealed when LER and dry weight were corrected for the effects
of other traits in the multivariate principal component analysis.
In summary, research has made progress in defining how
plant morphology criteria could be used in a plant improvement
context. However, the cost and logistics of making the
required trait measurements on a large number of plants are
formidable and there may also be trade-offs between desirable
traits.
Perennial ryegrass breeding in New Zealand Crop & Pasture Science 117

Environment
(light, water, nutrients, temperature)

LER LAR

LED
Site filling

Leaf
length

Potential
tiller bud
number
LAI

Tiller
weight

Tiller Tiller
appearance death

–3/2 Self-
thinning law

Tiller
Yield
density

PRODUCTIVITY PERSISTENCE

Fig. 4. Relationships between plant morphological characteristics (light grey boxes), persistence through
tiller population demography (dark grey boxes), and productivity (white boxes) (source: Bahmani 1999).
LER, Leaf elongation rate; LAR, leaf appearance rate; LED, leaf elongation duration; LAI, leaf area index.

Root morphology however, is not fully understood. In terms of development,

Larger, deeper rooting systems are likely to increase plant
water and nutrient uptake, potentially improving yield and
persistence of ryegrass. Experiments by Bonos et al. (2004)
and Crush et al. (2007, 2010) demonstrated that heritable
variation in root depth and weight and root system shape
exists, and associated QTLs were identified. Some of these
QTLs appear to have synteny with rice, meaning that findings
for rice may provide a ‘template’ for ryegrass. Although this
provides a starting point, there are still several questions
surrounding ‘ideal’ rooting system traits in ryegrass. Matthew
et al. (2001) calculated that while root hairs only make up 10%
of root mass, they make an important contribution to the total
root surface (~90%). Their involvement in nutrient uptake,
ryegrass roots are segmentally organised like leaves, taking
approximately five–six leaf appearance intervals to develop
(Matthew and Kemball 1997; Robin et al. 2010). The
development process is continual, and occurs simultaneously
on several adjacent phytomers (individual structural units that
in serial arrangement make up a tiller). Interestingly, in the
experiment of Robin et al. (2010), the length of the lateral
branches continued to increase after elongation of the main
root axis ceased. This is consistent with the hypothesis that
roots at the first root-bearing phytomers intercepted the
majority of photosynthate, with little reaching the older roots,
and that in substrate-starved older roots, continued peripheral
growth of some branches is driven by catabolism of other
118 Crop & Pasture Science
branches. Older roots may also be reliant on daughter tillers
for substrate supply, as was demonstrated in a tropical grass
(Carvalho et al. 2006); however, the root ageing process
in ryegrass is not well understood. Better understanding of the
root development and ageing process in ryegrass, and clarifying
which root morphological characteristics are functionally
important, will improve future selection for desirable root traits.
One important factor that contributes to reduced plant
persistence on some soil types, particularly peat, is pulling or
uprooting of ryegrass plants by cattle (Thom et al. 2003). Plants
that are more tolerant to pulling are likely to have one or more
of the following attributes: better anchorage through larger
root systems or longer roots (Crush et al. 2002), greater root
penetrating ability (Crush et al. 2002), and lower leaf shear
strength (associated with narrower leaves; Thom et al. 2003).
Variation exists in many root traits both within and between
cultivars (Crush et al. 2002, 2007); therefore, it should be
possible to breed plants that are more resistant to pulling.
Another factor that may contribute towards more pull-resistant
plants is increased aluminium (Al) tolerance. In many hill country
and upland soils in NZ, acidity and associated Al toxicity
are major constraints to root growth and, consequently, pasture
production (Edmeades and Wheeler 1985). Aluminium interacts
with root cells and/or their components to inhibit root elongation

(Ciamporová 2002; Haling et al. 2011), and as Al toxicity is more
likely in the subsoil than the topsoil, there is often poor root
penetration into the subsoil, leaving plants more susceptible to
pulling or death, particularly during dry conditions (Wheeler
and O’Connor 1998). Wheeler et al. (1992) identified ryegrass
plants with better Al tolerance than Grasslands Nui. When
grown in a reconstructed, acid soil profile, the plants with
greater Al tolerance had greater DM yields before and after
drought, and greater tiller survival than Grasslands Ariki,
Ellett, and Droughtmaster. The heritability of total yield in the
presence of Al averaged 0.33, indicating that although progress
could be made, it would likely be slow.
Herbage yield
It is important that the annual herbage yield of new perennial
ryegrass cultivars continues to improve; however, in the future,
greater emphasis may be placed on gains in certain seasons, and/
or cultivars that demonstrate slightly lower gains in annual
production but enhanced persistence through time. Chapman
et al. (2011b) estimated that in well-managed, pasture-based
dairy systems in southern Australia, additional DM was worth
AU$0.28–0.34/kg extra gross farm income if available in
summer, compared with $0.11–0.24/kg DM if available as a
pro-rata increase in every month of the year. This reflects the
seasonal supply of pasture feed relative to feed demand in dairy
regions of southern Australia. Chapman et al. (2011b) also
observed differences in the relative value of additional feed in
different seasons depending on location, and noted that the
economic value of additional feed will also differ when major
strategic management factors such as stocking rate and use of
purchased feed are taken into account. McEvoy et al. (2011)
developed an economic ranking index for perennial ryegrass
cultivars in Ireland to determine which traits had the greatest
impact on the economic performance of the farm system. Using a
J. M. Lee et al.
baseline scenario, spring and autumn DM yields were three to
five times more valuable than the midseason yield, respectively.
Although no published information is available, a similar index
is currently under development in NZ for perennial ryegrass
cultivars which takes into account the economic value of
additional feed at different times of the year in the major
dairying areas of NZ. As well as giving farmers information
on the relative ranking of perennial ryegrass cultivars according
to their traits and the estimated economic value of those traits, the
index system should explicitly identify the traits of greatest
economic importance and thereby assist in the development of
objectives for future plant-breeding programs.
Herbage nutritive value
Kolver and Muller (1998) demonstrated that cows fed a pasture-
only diet produced less milk than cows fed a balanced total mixed
ration (29.6 v. 44.1 kg/cow.day). The Cornell Net Carbohydrate
and Protein System identified DMI as a key factor influencing
milk production, with 61% of the reduction in milk production
explained by the lower DMI of the pasture-fed cows (19.0 v.
23.4 kg DM/cow.day). Any trait, therefore, that increases the
DMI of pasture-fed cows is likely to enhance milk production.
Bryant (2009) examined the effect of differences in chemical
components and digestion rate of different ryegrass lines using a
rumen-animal feeding decision model. In agreement with earlier
work by Kolver et al. (1998), the three factors with the greatest
impact on DMI and, therefore, milk production were low lignin
content, low fibre content, and fast rates of fibre degradation.
In ryegrass, the fibre components currently receiving most
attention are lignin and hydroxycinnamic acids such as diferulic
acids (Faville et al. 2010). Both compounds limit or obstruct
cell-wall degradation, and heritability estimates are high enough
in grasses to attempt breeding for these traits (Casler et al.
2008). Thus far, perennial ryegrass populations from diverse
origins have been analysed for fibre component concentration
and in sacco degradation kinetics. Degradation rates were
correlated with several fibre components, including certain
diferulic acids, diphenolic acids, and cell-wall oligomers,
indicating that selection of plants using these compounds may
enhance degradation rates (Faville et al. 2010). In addition, the
extensive genotypic variation in diferulic acids and other fibre
components among 70 diverse perennial ryegrass populations
indicates that useful sources for reducing the diferulic acid
concentrations exist (Faville et al. 2010). Further work is
under way to improve the fibre degradability of ryegrass.
Water deficit tolerance
Over the next four decades it is predicted that annual rainfall
is likely to decrease in NZ, resulting in most of the North Island
of NZ, and the Canterbury plains stretching into eastern
Southland, spending up to 10% more time in drought (NIWA
2011). This will place additional pressure on perennial ryegrass
growth and persistence. Breeding ryegrasses with increased
tolerance to soil water deficits should, therefore, become a
higher priority.
Plants have various strategies for adapting to water deficits
(Turner 1986; Ludlow 1989), namely drought avoidance,
dehydration postponement, or dehydration tolerance (Table 3).
Different morphological and physiological traits confer various
Perennial ryegrass breeding in New Zealand
Table 3. Mechanisms of plant adaptation to soil water deficit and their influence on plant productive processes as defined by
Turner (1986)
Mechanism
Drought avoidance
Rapid phenological developmentA
Developmental plasticity
Dehydration postponement
Maintenance of turgor
Maintenance of water uptake
Increased root density and/or depth
Increased liquid-phase conductance
Reduction of water loss
Reduction of leaf area
Increase in stomatal and cuticular resistance
Reduction in radiation absorbed
Osmotic adjustment
Maintenance of volume
Increase in elasticity
Dehydration tolerance
Protoplasmic tolerance
A
No correlation was found between early flowering and drought survival in ryegrass cultivars; however, the early flowering cultivar had
alpine origins (Volaire et al. 1998).
forms and degrees of tolerance, which means that identifying
a plant-breeding strategy is not straightforward. In addition,
perennial ryegrass is not considered to be well adapted to
water deficits (Jung et al. 1996). When water and nutrients are
sufficient, perennial ryegrass competes strongly for light with
other species by allocating a high proportion of growth resources
to shoot growth while maintaining a comparatively shallow root
system (Boschma and Scott 2000; Boschma et al. 2003a, 2003b).
In common with most of the higher plant species, the root mass
fraction in ryegrass increases as below-ground resources start
to limit growth (Cullen et al. 2006). However, when both
above- and below-ground resources simultaneously limit
growth, resource allocation patterns in ryegrass favour shoot
growth (Cullen et al. 2006). By contrast, phalaris (Phalaris
aquatica L.), which is more tolerant of soil moisture deficits
than perennial ryegrass (Boschma and Scott 2000), allocates a
higher proportion of growth resources to the root system in
response to simultaneous growth stresses (Cullen et al. 2006).
This difference in growth strategies between species indicates
that improving the tolerance of perennial ryegrass to water
deficit may be a difficult breeding objective. Nonetheless,
there is variation within the species in some key drought-
tolerance traits, indicating that progress is possible. For
example, perennial ryegrass genotypes demonstrate marked
variation in their ability to penetrate compacted areas and/
or root depth (Crush et al. 2002; Bonos et al. 2004), their
response to water deficit (i.e. increased or decreased root
growth; Crush et al. 2007), and their ability to recover from
water deficit stress (i.e. maintenance of live leaf area and tiller
survival following drought; Cheplick et al. 2000).
Nitrogen use efficiency
There is increasing concern worldwide as to the effect of intensive
farming on surface water quality, with nitrate leaching one of the
Crop & Pasture Science 119
Productive Additional references
processes reduced?
No Silsbury (1961); Humphreys et al. (2006)
No
No Chaves et al. (2003); Nie and Norton (2009);
Zhou et al. (2009)
No
Yes
Yes Butler (2008); Zhou et al. (2009)
Yes
No
No
Yes
main concerns. Perennial ryegrass has 80% of its root mass in the
top 15 cm of soil (Troughton 1951); therefore, excess nitrate
(predominantly from urine patches) only needs to leach a short
distance through the soil profile to become inaccessible to plants.
In the future, breeding objectives should focus on improving the
ability of perennial ryegrass to capture nitrate, thus reducing
the environmental impact of farming. This may include breeding
plants with greater winter growth (Crush et al. 2005; Popay and
Crush 2010), greater rooting depth (Crush et al. 2007; Popay and
Crush 2010), and larger (Crush et al. 2007) or more finely divided
root systems (Crush et al. 2005). Crush et al. (2010) demonstrated
that one cycle of selection for adventitious root system shape was
sufficient to create progeny with differing vertical distribution of
root mass, indicating that gains in this area using conventional
breeding are possible. Challenges for such research will be to
identify which root morphological traits are most beneficial and
methodologies to select for those traits.
Overseas, ryegrass breeding has developed lines with
divergent N-use efficiency, defined in these cases as either leaf
DM produced per unit of whole plant N uptake or greater plant
growth under limited N supply. In the UK, the ryegrass breeding
line with greater N-use efficiency had greater leaf regrowth rates
and reduced N concentration in the leaves (Wilkins et al. 1997,
2000). It also maintained a greater proportion of leaf in the sward
throughout the year, produced a third less reproductive tillers
during heading, and survived better over the 3-year experiment
than the line with lower N-use efficiency (Wilkins and Rognli
2002). Gains in digestibility from reduced flowering, however,
were not as great as expected; the annual DM yield was 7% less
than the control under simulated grazing, and the more N-use-
efficient selection could not be produced commercially due to
low seed yield. No further work, therefore, was conducted on this
selection (P. W. Wilkins, pers. comm.). In the Netherlands, plants
selected for increased LER in hydroponic culture at low N had
120 Crop & Pasture Science
higher herbage accumulation rates, higher tillering capacity, and
a higher leaf weight ratio (Van Loo 1993; Van Loo et al. 2003).
These plants yielded more in the field under low N supply.
Unfortunately, it is not known whether these selections were
subsequently used in cultivar development (E. N. Van Loo, pers.
comm.).
Disease and pest resistance
It is important that the disease resistance of new perennial
ryegrass cultivars continues to improve, with rust-susceptible
lines eliminated during the cultivar development and
evaluation process. Endophyte is likely to continue to be a key
plant-breeding focus, as recognised by Easton (2007), who
concluded that the endophyte status of a perennial ryegrass
cultivar is vitally important to its value, often more so than the
particular cultivar itself. Grass grub remains the only major NZ
insect pest against which current perennial ryegrass endophytes
have little activity. Endophytes found in tall fescue and meadow
fescue that produce loline alkaloids show some promise for
protection against grass grub (Popay et al. 2003; Patchett et al.
2008); however, the lack of stable transmission of these loline-
producing endophytes in perennial ryegrass seed is a major
obstacle to overcome (Easton et al. 2009a). This is an area of
active research. Identification of novel endophytes covering the
spectrum of insect pests (and potential new pests) will remain vital
in future plant breeding. Improving novel endophyte transmission
rate through seed generations, and better endophyte longevity
in stored seed, also remain challenges to delivering seed with high
endophyte levels to farmers.
Integrating breeding, agronomy, and farming systems:
a strategy for the future
New Zealand pastures are complex, with most containing not only
ryegrass (often more than one genotype), but also an associated
endophyte strain, white clover, and other plant species. Although
there is evidence that genetic gains in the herbage yield of
perennial ryegrass have been made through plant breeding
over the past four decades, it is not clear how much gain has
been achieved in pasture performance as a result of this progress,
or whether this gain is sufficient to secure the competitiveness
of pasture-based dairy systems into the future. To address these
questions, the disciplines of plant breeding, pasture agronomy/
ecology, and farm systems analysis must be more closely
integrated than has been the case in the past. Here, we propose
three areas where new knowledge is required to fully understand
the value that is being delivered to dairy farm businesses through
perennial ryegrass breeding. Many other areas could, and should,
be identified; however, these three at least illustrate how different
disciplines could contribute.
First, economic benefit to dairy farm businesses of different
productivity-related traits in perennial ryegrass should be
determined and communicated to farmers and plant breeders.
Using agronomic information and farm system simulation tools,
it is possible to estimate the gross economic return to farming
businesses of incremental changes in traits such as seasonal
DM production (Brookes et al. 1993; Chapman et al. 2011b)
or nutritive value (Bryant 2009). McEvoy et al. (2011) integrated
such information into a weighted index that can be used to rank
J. M. Lee et al.
ryegrass cultivars according to likely economic value to a farm
business, akin to the Breeding Worth and Production Worth
indices used to rank dairy sires in the NZ animal evaluation
system (Harris et al. 1996). There is sufficient information in
such analyses to direct plant-breeding effort into the traits that
have the greatest impact on estimated economic value. There
are, however, technical hurdles to overcome, not least being the
availability of reliable and comprehensive data on the key traits
for all commercially available material. There is a substantial
store of DM yield information from NFVT conducted over the
past two decades, which provides a starting point for economic
analysis. However, data on persistence and nutritive value are less
plentiful. In addition, well-targeted evaluation procedures will be
needed if such information is to be generated routinely and with
sufficient reliability for the development of robust economic
ranking indices for the NZ dairy industry.
Second, the relationships between perennial ryegrass traits
and whole pasture performance should be defined and
accommodated in the development of quantitative forage value
indices. While there is a substantial store of NFVT yield data
available for perennial ryegrass cultivars, it comes predominantly
from monoculture plots under high N supply, whereas farmers
generally manage mixed ryegrass/clover pastures. The translation
of information on cultivar-specific traits to pasture performance
measures needs to be addressed, including the possibility that
cultivar rankings do not necessarily equal pasture performance
rankings, and/or that the range between poorest and best cultivars
may differ when rankings from ryegrass monocultures are
compared with rankings for mixed pasture (i.e. Eerens et al.
2001, their Table 2, where the increase in ryegrass yield from
late 1990s ryegrasses compared with 1980s ryegrasses was 4–7%
greater than the increase in total pasture yield). Both relative
and absolute economic values associated with cultivars at the
whole-pasture level are important. Again, it appears that well-
targeted agronomic research is required to address this specific
issue by building upon the existing comprehensive store of
knowledge regarding inter-specific competition in pastures,
especially grass–clover–endophyte interactions (e.g. Chapman
et al. 1996; Schwinning and Parsons 1996).
Third, G  E interactions should be better documented for the
dairying regions of NZ. It is noted here that it is necessary to
include both endophyte strain and plant host under the definition
of ‘genotype’, and ‘environment’ must include the management
environment (i.e. effects of management), as well as the physical
macro-environment. The issue of interactions between perennial
ryegrass and white clover noted above is a case in point. In
environments which favour ryegrass dominance, such as those
receiving high rainfall/irrigation and high N-fertiliser inputs, a
close match between economic value rankings based on ryegrass
monoculture evaluation data and pasture performance could be
expected. Conversely, the relationship may be weak in low-N
environments subject to regular soil water stress which allows
other species to compete successfully in the pasture community.
Furthermore, the defoliation management applied could result in
the re-ranking of cultivars (Gilliland and Mann 2000; Orr et al.
2001; McEvoy et al. 2009), although there are no published
experiments for NZ material. These possibilities could have
significant implications for objectives for plant-breeding
programs and for the way cultivars are evaluated. To our
Perennial ryegrass breeding in New Zealand
knowledge, there has been no systematic agronomic investigation
of G  E interactions in the key productivity traits of modern
perennial ryegrass cultivars in NZ. Future progress in ryegrass
plant breeding and in the uptake of new cultivars by farmers may
be restricted if the issue is not addressed, and the uncertainty
regarding cause and effect that was evident, for example, in the
Proceedings of the Pasture Persistence Symposium (Mercer
2011), will continue. The return to the national economy from
public and private sector investment in ryegrass plant breeding is
likely to be suboptimal under these conditions.
Acknowledgments
The authors acknowledge the significant contribution of the anonymous
reviewers whose comments were highly valuable. Sincere thanks also to
Dr Alan Stewart and Michael Norriss (PGG Wrightson Seeds), Professor
Warren Williams (AgResearch) and Graham Kerr (New Zealand Agriseeds
Ltd) for their comments on the manuscript. Thanks to Dr H. S. Easton,
Dr P. W. Wilkins, and Dr E. N. Van Loo for clarification on past research. This
work was funded by New Zealand dairy farmers through DairyNZ.
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