1. Introduction
Perennial ryegrass (PRG,
Lolium perenne L.) is an important forage species grown on many dairy farms in south-eastern Australia, New Zealand, Ireland, the United Kingdom and continental Europe, with many cultivars of the species commercially available that have been developed over several decades of plant breeding. Early plant breeding initiatives concentrated on annual dry matter (DM) yield and plant survival. More recently, factors such as seasonal DM yield and nutritive quality have been addressed by plant breeders [
1]. This focus has led to the development of cultivars with a variety of flowering dates. Additionally, cultivars with an increased number of chromosomes (tetraploids) have been developed from diploid PRG as a strategy to increase nutritive value [
2]. Commercially available cultivars are now routinely categorised according to their ploidy (diploids or tetraploids) and maturity (heading and flowering) dates, with these characteristics commonly referred to as functional types [
3]. Tetraploids were widely used in Europe and the United Kingdom before being adopted in Australia [
2], offering a possible explanation for the relatively limited research investigating differences in nutritive characteristics between ploidy type on a seasonal basis and between individual cultivars applicable to Australian dairy environments. The Australian dairy industry is concentrated in the south-eastern states, with Victoria and Tasmania collectively producing 75% of Australia’s milk [
4]. Across the dairy regions in these states alone, key environmental elements such as rainfall vary from less than 500 mm to more than 1100 mm annually, with farms in lower rainfall areas requiring irrigation to support PRG pastures. Research in New Zealand has demonstrated differences between early and late maturing diploids and between diploids and tetraploids for metabolisable energy (ME), a key nutritive characteristic trait, with the magnitude of these differences varying between some seasons and in contrasting dairy environments [
5]. A recent study in Australia [
3] also found differences between PRG functional types but was limited to reporting results from one dairy environment.
Since the establishment of forage value indices in Ireland, New Zealand and Australia, there has been substantial interest in cultivar evaluation data for traits that are important on dairy farms. The Pasture Profit Index (PPI) in Ireland [
6,
7] was the first of these systems to be established and publishes individual cultivar information on seasonal yield, mid-season quality (digestibility), silage yield and persistence with these traits weighted according to their importance. In this system, the weighting for pasture digestibility is 25% [
8]. For the published cultivars, tetraploids on average have dry matter digestibility (DMD) values 6.7 and 7.5 g/kg DM higher than diploids for intermediate and late heading dates, respectively. These differences in DMD are equivalent to 0.11 and 0.13 megajoules (MJ) of ME per kg of DM. The New Zealand Forage Value Index (FVI) [
9] has progressively added new traits (ME and persistence) since the first release which ranked cultivars based on the value of seasonal DM yield. It publishes differences in ME between functional groups for two unique environments on a seasonal basis, suggesting a lack of sufficient cultivar-specific data for this trait. In the upper North Island, tetraploids have seasonal ME values 0.23–0.46 and 0.24–0.35 MJ/kg DM greater than mid- and late-heading diploids, respectively. For the rest of New Zealand, based on trial data from the Canterbury region in the upper South Island [
10], tetraploids have seasonal ME values 0.42–0.64 and 0.27–0.35 MJ/kg DM greater than mid- and late-heading diploids, respectively. In contrast, the PRG Forage Value Index (FVI) developed in Australia is currently limited to providing information on seasonal dry matter production due to the lack of individual cultivar information on nutritive characteristics relevant to Australian dairy environments [
11,
12]. In New Zealand, ME concentration is considered an “indicator trait” for nutritive value in the FVI [
13], although it is recognised by these researchers that measures of other nutritive characteristics (neutral detergent fibre, NDF and crude protein, CP) should be used in conjunction with ME to comprehensively evaluate the feeding value of forages [
13,
14].
Phenotypic information on these three traits (ME, CP and NDF) measured on modern cultivars in contrasting dairy environments is therefore needed, as is investigation of the seasonal variability in these traits. Collecting data on nutritive characteristics is labour-intensive and costly. Therefore, a validation of the range and variability of nutritive characteristics and a comparison of the ranking of cultivars at different locations based on their nutritive characteristics are warranted prior to the collection of a more comprehensive data set.
The study reported aimed to (i) investigate the variation in key nutritive characteristics: ME, crude protein (CP) and neutral detergent fibre (NDF) of diploid and tetraploid perennial ryegrass cultivars at four trial sites located in four south-eastern Australian dairy regions at multiple harvests over a 3-year period and (ii) compare the ranking of cultivars between sites and seasons for these characteristics. These results provide information on the amount of variation in nutritive characteristics in modern cultivars across a range of Australian dairy environments. The results will be of benefit to breeding companies, where improved nutritive characteristics have been identified as a target, and to dairy farmers when choosing cultivars to sow. One of the key tools that farmers use to choose varieties is the FVI, and the main impediment to including nutritive characteristics in the FVI is the lack of data. These results illustrate the value of including nutritive characteristics in the FVI and describe a practical methodology to address this lack of data.
4. Discussion
This study will facilitate the expansion of the Australian FVI to include nutritive traits of PRG that are economically important in pasture-based dairy production systems and more broadly add to the body of literature on the variation in nutritive characteristics of PRG cultivars used in temperate dairy systems. The lack of any measure of nutritive value in the Australian FVI was identified as a limitation of the index in its development phase [
11]. However, the lack of data on the nutritive characteristics of modern PRG cultivars in south-eastern Australian dairy environments precluded the inclusion of this information at that time. In contrast, the PPI in Ireland [
6,
7] and more recently the DairyNZ FVI [
9] have included nutritive characteristic traits. However, the New Zealand system is currently limited to reporting ME at the functional group level rather than for individual cultivars, further indicating the paucity of information available on the nutritive characteristics of modern individual PRG cultivars in the temperate dairying areas of the southern hemisphere.
Metabolisable energy is the primary determinant of milk production [
21] and therefore is considered an economically important nutritive characteristic trait. It is widely known that the ME of pastures fluctuates seasonally, and studies have shown that tetraploid PRGs have higher digestibility and ME values than diploid PRGs [
3,
22,
23]. In Australia and New Zealand, most tetraploids are classified as “late” or “very late” heading, enabling them to produce more vegetative growth in late spring [
3] compared to some of the diploid cultivars that produce seed heads earlier in spring. Late heading can therefore lead to pastures with superior nutritive characteristics in spring when PRG growth is at its peak. The heading dates of diploids range from early to late, but even the late-heading diploid cultivars have been found to have a lower ME concentration than comparable tetraploids [
3]. Tetraploid cultivars have larger cells than diploids, leading to differences in digestibility [
2]. However, it is important to note that there is variation amongst cultivars with some diploid candidates in our study having equivalent ME concentrations to some tetraploid candidates in all seasons of the year, even though on average the tetraploid candidates had a higher ME.
Trial design and the method of statistical analysis used can affect whether differences between cultivars for traits of interest are detected [
24]. Our analyses accounted for the spatial and repeated measurement structure of the trial data, which was one of the ways suggested [
24] to improve the detection of candidate differences. The power analysis of ME was conducted on the average performance of candidates across seasons at each trial site and showed that four replications were adequate for the detection of a 0.3 MJ/kg DM ME difference between candidates. The Ellinbank trial had lower power to detect this change than the other sites mainly due to ME measurements in Ellinbank being more variable than the other sites. Candidate performance varied by season at each site. While it would be possible to perform a seasonally specific power analysis for each site, the result will be a different number of replications needed to achieve a certain power for each season. This could not be practically applied as it is not possible to change the number of replicates each season.
It is never possible to test all cultivars in all environments as even within regions, there is variation in environmental factors such as rainfall and soil type. However, the results comparing candidate rankings for ME from this study indicate that it is possible to use trial data from other dairying areas of south-eastern Australia where necessary for the purpose of ranking cultivars on ME. A multi-environment approach, as is currently done with the DM yield data in the Australian FVI, could also be considered for this trait. Our recommended strategy for the testing of ME across south-eastern Australian dairy environments is to have every cultivar tested in at least one fully replicated trial where all plots are tested individually on each sampling occasion to achieve the greatest accuracy and precision [
16], which could then be supported by a network of trials that may adopt the composite sampling strategy (if resources are constrained) to achieve geographic breadth. As the knowledge of the environments in the trial network grows, it would become possible to allocate the fully replicated trials to those environments that are inherently more variable. Farmer confidence in the FVI is likely to be greater if they know that cultivars have been tested in their region.
The results for CP were less conclusive, with trends in CP varying between the trial sites and inconsistent trends, and in some cases, no differences between functional groups were observed. CP is currently not included in either the PPI or the DairyNZ FVI as the CP content of PRG is not considered a limiting nutrient for dairy cows grazing these pastures [
25].
In this study, the NDF of most tetraploids was consistently lower (mean across seasons and sites 45.3% DM) than the diploid cultivars (mean across seasons and sites 46.8% DM) throughout the year, which was expected as NDF is inversely related to digestibility and ME. While high ME and low NDF forages may be desirable, there are on-farm pasture and feed management considerations associated with this. Both managing grazing residuals to balance pasture regrowth and utilisation and consideration of the overall NDF concentration of dairy cow diets are important. Recent Irish research found lower post-grazing residuals in tetraploids (3.7 cm) compared to diploids (4.1cm), with the lower residuals in this context associated with a higher grazing efficiency [
26]. However, low post-grazing residuals can affect pasture regrowth [
27]. Notably, the residuals to achieve high grazing efficiency in Ireland were 26% lower than recommendations in Australian and New Zealand dairy systems [
27,
28], where a residual grazing height of 5 cm is considered optimal for growth and persistence [
29]. In Australian dairy systems, cereal grain supplements low in NDF are commonly fed to cows. The NDF values of these feeds must also be considered when selecting forages in these systems to ensure that the NDF content of cow diets is within the recommended range of 30–40% of DM intake, which is considered sufficient to support rumen function but not limiting total intake. [
21].
5. Conclusions
Our study showed that the tetraploid PRGs had consistently higher ME values than diploid PRGs, although the differences were numerically small and there was evidence of variation within diploid and tetraploid groups. These data show that it would be possible for dairy farmers to select cultivars with above-average nutritive characteristics in their environment. In some cases, candidates with the higher ME values were also high yielding candidates that would increase the on-farm benefits of sowing cultivars with these characteristics. High yielding cultivars with high ME may also provide sources of elite germplasm for breeding programs. The inclusion of an economic value for marginal differences in ME may result in greater differentiation between the seasonal performance of cultivars when this trait is added to the Australian FVI. Ascertaining whether this translates into tangible on-farm milk production benefits and profit would require complementary farm systems studies and/or modelling.
The ranking of candidates for ME was more consistent across trial sites compared to CP and NDF, suggesting these two traits, especially CP, are more sensitive to environmental influences. Further research with a larger dataset would enable a greater understanding of the genotype by environmental interactions for these traits.