This review examines the use of changes in soluble carbohydrate reserves, and the onset of senescence in ryegrass (Lolium spp.), as key criteria for successfully managing an intermittent grazing system for dairy cattle. Ryegrass is a ‘3-leaf ’ plant; that is, only about 3 green leaves/tiller exist at any one time with the initiation of a new leaf coinciding with senescence of the oldest fourth leaf. Thus, grazing pasture older than 3 leaves/tiller will not only lead to wastage of pasture but also the senescent material will reduce overall quality of herbage. Based on this, the time taken for 3 new leaves/tiller to regrow sets the maximum grazing interval. On the other hand, in a well-utilised dairy pasture, most ryegrass leaf has been removed and the plant relies on stored water-soluble carbohydrate reserves to grow new shoots and hence regain photosynthetic capacity. If the concentration of water-soluble carbohydrates is inadequate, because there has been insufficient time to replenish in the previous inter-grazing period, regrowth will be suppressed and this may also affect persistence in the longer term. Immediately after grazing, water-soluble carbohydrate reserves decline as they are used to regrow new shoots, and root growth stops. It is not until about 3/4 of a new leaf/tiller has regrown that the plant has adequate photosynthetic capacity for growth and maintenance and only then does water-soluble carbohydrate replenishment and root growth commence. Studies have shown that subsequent regrowth is suppressed if plants are redefoliated before the 2 leaves/tiller stage of regrowth. Also, the levels of potassium and nitrogen (as nitrates and other non-protein nitrogen products) may be very high and cause metabolic problems in stock grazing such pasture. Thus, replenishment of water-soluble carbohydrate reserves sets the minimum grazing interval at 2 leaves/tiller. The rate of accumulation of water-soluble carbohydrates in the plant is a function of input through photosynthesis (source) and output to growth and respiration (sinks). Thus, apart from grazing interval (which sets the time to replenish water-soluble carbohydrate plant reserves), water-soluble carbohydrate storage will be influenced by incoming solar radiation (cloud cover, day length, pasture canopy density) and energy needs of the plant through respiration (temperature, canopy mass) and growth. Relating grazing interval to leaf number places the emphasis on the readiness of plants to be grazed rather than on the animals’ requirements, with leaf appearance interval depending primarily on ambient temperature. This allows grazing interval to be expressed in a similar morphological stage of growth, irrespective of season or location. Setting grazing interval on these 2 criteria has been shown to maximise growth and persistence of ryegrass and optimise the levels of most nutrients in pasture required by dairy cattle including protein, water-soluble carbohydrates, calcium, potassium and magnesium. Metabolisable energy and fibre do not change appreciably up to the 3 leaves/tiller stage of regrowth. On the other hand, grazing pasture before 2 leaves/tiller not only retards regrowth and reduces persistence, it provides forage too high in potassium and protein (nitrates) and too low in water-soluble carbohydrates for dairy cattle.
A glasshouse study was undertaken to determine the priority within the perennial ryegrass (Lolium perenne L.) plant for leaf and root growth and daughter tiller initiation after defoliation, in relation to various levels of water‐soluble carbohydrate (WSC) reserves at defoliation. Individual plants were arranged in mini‐swards, and underwent varying defoliation frequencies and ambient temperatures before defoliation, and harvest heights at defoliation, to obtain a gradient of WSC content at H1, the date when all plants were defoliated. Defoliation interval consisted of defoliating either three times at the one new leaf tiller–1 stage (1‐leaf stage) of regrowth, or once only at the 3‐leaf stage, up to H1, while night temperature in the week prior to H1 was altered from 15°C to either 8 or 20°C. At H1, plants were defoliated to a stubble height of either 20 or 50 mm. Plants were subsequently destructively harvested at days 4, 6, 8, 12, 18 and 27. Leaf and root extension and tiller dynamics were also measured. On a regrowth timescale, tiller initiation was most sensitive, root regrowth moderately sensitive, and leaf regrowth relatively insensitive to a decrease in WSC. The time of daughter tiller initiation also coincided with replenishment of stubble WSC levels. In contrast to this sequence of regrowth events following defoliation, the quantitative effects on growth were different, with elongation and survival of roots most affected by reduced WSC levels. A 30‐fold difference in stubble WSC at H1 between high and low WSC plants (1·52 vs. 0·05 mg tiller–1) produced only a 4‐fold increase in leaf dry matter (DM) after 27 d (2·2 vs. 0·6 g plant−1), while tiller number plant−1 increased 6‐fold (138 vs. 23% increase in tiller number from H1). Root elongation rate was 59 times higher in the high than in the low WSC plants (1·18 vs. 0·02 mm d−1). From a pasture management perspective, the study confirms that defoliation, coinciding with the 3‐leaf stage of regrowth and around a stubble height of 50 mm, optimizes persistence and productivity of perennial ryegrass. By allowing more rapid replenishment of WSC reserves, this optimal defoliation strategy enables a greater proportion of WSC to be allocated to maintain a more active root system, and promotes tillering, compared with more frequent and close defoliation.
BackgroundPerennial ryegrass (Lolium perenne L.) is an important pasture and turf crop. Biotechniques such as gene expression studies are being employed to improve traits in this temperate grass. Quantitative reverse transcription-polymerase chain reaction (qRT-PCR) is among the best methods available for determining changes in gene expression. Before analysis of target gene expression, it is essential to select an appropriate normalisation strategy to control for non-specific variation between samples. Reference genes that have stable expression at different biological and physiological states can be effectively used for normalisation; however, their expression stability must be validated before use.ResultsExisting Serial Analysis of Gene Expression data were queried to identify six moderately expressed genes that had relatively stable gene expression throughout the year. These six candidate reference genes (eukaryotic elongation factor 1 alpha, eEF1A; TAT-binding protein homolog 1, TBP-1; eukaryotic translation initiation factor 4 alpha, eIF4A; YT521-B-like protein family protein, YT521-B; histone 3, H3; ubiquitin-conjugating enzyme, E2) were validated for qRT-PCR normalisation in 442 diverse perennial ryegrass (Lolium perenne L.) samples sourced from field- and laboratory-grown plants under a wide range of experimental conditions. Eukaryotic EF1A is encoded by members of a multigene family exhibiting differential expression and necessitated the expression analysis of different eEF1A encoding genes; a highly expressed eEF1A (h), a moderately, but stably expressed eEF1A (s), and combined expression of multigene eEF1A (m). NormFinder identified eEF1A (s) and YT521-B as the best combination of two genes for normalisation of gene expression data in perennial ryegrass following different defoliation management in the field.ConclusionsThis study is unique in the magnitude of samples tested with the inclusion of numerous field-grown samples, helping pave the way to conduct gene expression studies in perennial biomass crops under field-conditions. From our study several stably expressed reference genes have been validated. This provides useful candidates for reference gene selection in perennial ryegrass under conditions other than those tested here.
Effect of defoliation management, based on leaf stage, on perennial ryegrass (Lolium perenne L.), prairie grass (Bromus willdenowii Kunth.) and cocksfoot (Dactylis glomerata L.) under dryland conditions. 1. Regrowth, tillering and water-soluble carbohydrate concentration Abstract A field study was undertaken between April 2003 and May 2004 in southern Tasmania, Australia to quantify and compare changes in herbage productivity and water-soluble carbohydrate (WSC) concentration of perennial ryegrass (Lolium perenne L.), prairie grass (Bromus willdenowii Kunth.) and cocksfoot (Dactylis glomerata L.) under a defoliation regime based on leaf regrowth stage. Defoliation interval was based on the time taken for two, three or four leaves per tiller to fully expand. Dry-matter (DM) production and botanical composition were measured at every defoliation event; plant density, DM production per tiller, tiller numbers per plant and WSC concentration were measured bimonthly; and tiller initiation and death rates were monitored every 3 weeks. Species and defoliation interval had a significant effect (P < 0AE05) on seasonal DM production. Prairie grass produced significantly more (P < 0AE001) DM than cocksfoot and ryegrass (5AE7 vs. 4AE1 and 4AE3 t DM ha )1 respectively). Plants defoliated at the two-leaf stage of regrowth produced significantly less DM than plants defoliated at the three-and four-leaf stages, irrespective of species. Defoliation interval had no effect on plant persistence of any species during the first year of establishment, as measured by plant density and tiller number. However, more frequent defoliation was detrimental to the productivity of all species, most likely because of decreased WSC reserves. Results from this study confirmed that to maximize rates of regrowth, the recommended defoliation interval for prairie grass and cocksfoot is the four-leaf stage, and for perennial ryegrass between the two and three-leaf stages.
Pasture Management 122A grono my J our n al • Volu me 10 0 , I s sue 1 • 2 0 0 8
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