Pastoral-based animal production systems are under increasing pressure to provide the high quantity and quality of feed needed for optimal ruminant performance. The capacity of farmers to increase forage yield further, solely by increasing fertilizer inputs or through improved pasture management, is limited. Emerging requirements to balance industry production targets against the need to reduce greenhouse gas emissions and N losses pose further challenges. Plant breeding is being asked to deliver results more urgently than at any time previously, and this review attempts to highlight issues that might limit the prospects for future progress by seeking lessons from four past examples: (i) white clover breeding gains and the need to consider the complexity of the grazed grass-clover mixed sward, with its tendency for cycling in plant species composition; (ii) a systems field trial of new and old grass ⁄ clover cultivars, and how the complexity of growth of perennial forage crops, and the dynamic optimality required for sustainable harvesting might limit our ability to breed for 'yield' per se; (iii) the manipulation of a physiological trait (low 'maintenance' respiration) and the implications of such changes for plant fitness and G · E interactions; and (iv) an hypothesis-driven development of a trait (high-sugar grasses) and the value of 'proof of concept' studies, the requirement of scientific understanding of the mechanisms of trait expression, and how one might in future go about assessing breeding achievements. We discuss the general ecological considerations around shifts in the frequency distribution of traits in new populations, whether altered conventionally or by genetic modification, and how selection for a particular trait might inadvertently reduce both fitness and persistence. A major priority for breeding, we propose, might be to revisit previously abandoned traits that affected the physiological performance of forage species, armed now with a capacity to monitor gene expression at the molecular level, and so unravel ⁄ control the G · E interactions that limited their benefits. We also discuss how a 'loss of yield advantage' of new cultivars, seen when tested several years after sowing, requires urgent investigation and propose this might be associated with fitness costs of perenniality. Finally, we argue for a careful reconsideration of what are realistic expectations for systems field trials and that focus on forage breeding might be shifted more to 'proof of concept' studies, critical experimental design, comparing 'traits' rather than 'cultivars', and the wider ecological assessment of fitness and function of traits in the plant, community and ecosystem.

Effects of elevated CO2 (700/.tL L -l) and a control (350/tL L -I CO2) on the productivity of a 3-year-old ryegrass/white clover pasture, and on soil biochemical properties, were investigated with turves of a Typic Endoaquept soil in growth chambers. Temperature treatments corresponding to average winter, spring, and summer conditions in the field were applied consecutively to all of the turves. An additional treatment, at 700/IL L-i CO2 and a temperature 6°C higher throughout than in the other treatments, was included.Under the same temperature conditions, overall herbage yields in the '700 #L L-l CO2' treatment were ca. 7% greater than in the control at the end of the 'summer' period. Root mass (to ca 25 cm depth) in the '700 juL L-CO2' treatment was then about 50% greater than in the control, but in the '700 ~L L -1 CO2 + 6°C ' treatment it was 6% lower than in the control. Based on decomposition results, herbage from the '700 ~tL L-l + 6oc , treatment probably contained the highest proportion of readily decomposable components.Elevated CO2 had no consistent effect on soil total C and N, microbial C and N, or extractable C concentrations in any of the treatments. Under the same temperature conditions, it did, however, enhance soil respiration (CO2-C production) and invertase activity. The effects of elevated CO2 on rates of net N mineralization were less distinct, and the apparent availability of N for the sward was not affected. Under elevated CO2, soil in the higher-temperature treatment had a higher microbial C:N ratio; it also had a greater potential to degrade plant materials.Data interpretation was complicated by soil spatial variability and the moderately high background levels of organic matter and biochemical properties that are typical of New Zealand pasture soils. More rapid cycling of C under CO2 enrichment is, nevertheless, indicated. Futher long-term experiments are required to determine the overall effect of elevated CO2 on the soil C balance.

This study describes the successive stages of development of branches from axillary buds in fully rooted plants of Trifolium repens grown in near optimal conditions, and the way in which this developmental pathway differs when nodal root formation is prevented as plants grow out from a rooted base. Cuttings of a single genotype were established in a glasshouse with nodal root systems on the two basal phytomers and grown on so that nodal rooting was either permitted (+R) or prevented (-R). In +R plants, axillary tissues could be assigned to one of four developmental categories: unemerged buds, emerged buds, unbranched lateral branches or secondarily branched lateral branches. In -R plants, branch development was retarded, with the retardation becoming increasingly pronounced as the number of -R phytomers on the primary stolon increased. Retarded elongation of the internodes of lateral shoots on -R plants resulted in the formation of a distinct fifth developmental category: short shoots (defined as branches with two or more leaves but with mean internode length equal to, or less than, 10% of that of the immediately proximal internode on the parent stolon) which had reduced phytomer appearance rates but retained the potential to develop into lateral branches. Transfer of +R plants to -R conditions, and vice versa, after 66 d demonstrated that subsequent branch development was wholly under the control of the youngest nodal root present, regardless of the age and number of root systems proximal to it.

Two manipulative experiments tested hypotheses pertaining to the correlative control exerted by nodal roots on branch development of the distal non-rooted portion of Trifolium repens growing clonally under near-optimal conditions. The two experiments, differing in their pattern of excision to manipulate the number of branches formed at the first 9-10 phytomers distal to the youngest nodal root, each found that after 20 phytomers of growth the total number of lateral branches formed on the primary stolon remained between five and seven regardless of where the branches formed along the stolon. Additional treatments established that nodal roots influenced branch development via relationships among shoot sinks for the root-supplied resources rather than through variation in the supply of such resources induced by fluctuations in photosynthate supply to roots from branches. Regression analysis of data pooled from treatments of both experiments confirmed that shoot-sink relationships for root- supplied resources controlled the branching processes on the non-rooted portion of plants. A disbudding treatment, which removed all the apical and axillary buds present on basal branches, but left other branch tissues intact, increased branch development of the apical region in the same way as did complete excision of the basal lateral branches. The apical buds and the elongation processes occurring immediately proximal to the buds were thus identified as strong sinks for the root-supplied resources. Such results suggest that branch development on the non-rooted shoot portion distal to the youngest nodal root is regulated by competition among sinks for root-derived resources, of limited availability, necessary for the processes of elongation of axillary buds and the primary stolon apical bud.

This chapter focuses on the influence of defoliation strategy on tissue flows and primary production in grasslands, and the scope for manipulating the balance between tissue growth and loss, soil C and N resources, and the potential impacts of climate change on the sustainability of grassland systems. It also discusses the principles of growth and utilization of grass and the effects of pasture utilization on the uptake, cycling and fate of C and N.