Kevin Foster

Genetic improvement of subterranean clover (Trifolium subterraneum L.). 1. Germplasm, traits and future prospects

Abstract. Subterranean clover (Trifolium subterraneum L.) is the most widely sown annual pasture legume species in southern Australia, valued in the livestock and grains industries as a source of high-quality forage and for its ability to fix atmospheric nitrogen. From its initial accidental introduction into Australia in the 19th Century and subsequent commercialisation in the early 1900s, 45 cultivars have been registered in Australia. These consist of 32 cultivars of ssp. subterraneum, eight of ssp. yanninicum, and five of ssp. brachycalycinum and range in flowering time from 77 to 163 days from sowing, enabling the species to be grown in a diversity of rainfall environments, soil types, and farming systems. Eleven of these cultivars are introductions from the Mediterranean region, 15 are naturalised strains collected in Australia, 18 are the products of crossbreeding, and one is derived from mutagenesis. Cultivars developed in Italy have been commercialised for the local market, whereas other cultivars developed in Spain, Portugal, and France have not had commercial seed production. Important traits exploited include: (i) selection for low levels of the oestrogenic isoflavone formononetin, which causes reduced ewe fertility; (ii) increased levels of dormancy imposed by seed-coat impermeability (hard seeds) for cultivars aimed at crop rotations or unreliable rainfall environments; (iii) strong burr-burial ability to maximise seed production; (iv) resistance to important disease pathogens for cultivars aimed at medium- and high-rainfall environments, particularly to Kabatiella caulivora and root rot pathogens; (v) resistance to pests, particularly redlegged earth mites; and (vi) selection for unique leaf markings and other morphological traits (where possible) to aid cultivar identification. Cultivar development has been aided by a large genetic resource of ∼10 000 accessions, assembled from its centre of origin in the Mediterranean Basin, West Asia, and the Atlantic coast of Western Europe, in addition to naturalised strains collected in Australia. The development of a core collection of 97 accessions, representing almost 80% of the genetic diversity of the species, and a genetic map, provides a platform for development of future cultivars with new traits to benefit the livestock and grains industries. New traits being examined include increased phosphorous-use efficiency and reduced methane emissions from grazing ruminant livestock. Economic analyses indicate that future trait development should focus on traits contributing to increased persistence and autumn–winter productivity, while other potential traits include increased nutritive value (particularly of senesced material), increased N2 fixation ability, and tolerance to cheap herbicides. Beneficial compounds for animal and human health may also be present within the species for exploitation.

Drought resistance at the seedling stage in the promising fodder plant tedera (Bituminaria bituminosa var. albomarginata)

Abstract. The perennial legume Bituminaria bituminosa (L.) C.H. Stirt. var. albomarginata (tedera) has been identified as a promising fodder plant for the southern Australian wheatbelt, but little is known about its drought resistance as a seedling. This study was conducted to (i) examine physiological and morphological responses to water stress of seedlings of tedera, in comparison with lucerne (Medicago sativa L.), biserrula (Biserrula pelecinus L.) and Afghan melon (Citrullus lanatus Thunb.), and (ii) investigate drought adaptation mechanisms of tedera seedlings. Seedlings were grown in a reconstructed field soil profile in pots in a glasshouse. By 25 days after sowing (DAS), plants of all species in the drought-stressed (DS) treatment had experienced water stress, with an average leaf relative water content (RWC) of 66% in DS compared with 79% in well-watered (WW) plants. Tedera, biserrula and Afghan melon maintained a higher RWC than lucerne. At 25 DAS, reductions in shoot dry matter in the DS treatment differed between species: 52% for Afghan melon, 36% for biserrula, 27% for lucerne, and no significant reduction for tedera. Paraheliotropic leaf angles of biserrula, lucerne and tedera were all higher in the DS treatment than in the WW treatment at 25, 32 and 52 DAS. This study revealed significant differences in rooting depth and stomatal conductance between the three legume species when under water stress, with tedera being the most drought-resistant. Traits that may allow tedera to survive a dry period following opening rains include vigorous seedling growth, early taproot elongation, effective stomatal control and paraheliotropic leaf movements.

Seasonal and diurnal variation in the stomatal conductance and paraheliotropism of tedera (Bituminaria bituminosa var. albomarginata) in the field

The mechanisms of drought resistance in perennial legumes are poorly understood. We explored the diurnal and seasonal variation (May, August, February) in stomatal conductance (gs) and paraheliotropism of three tedera accessions (Bituminaria bituminosa (L.) C.H. Stirton var. albomarginata) and lucerne (Medicago sativa L.), both perennial legumes, grown in the field. For the tedera accessions, there was a significant reduction in gs during the day in May (late autumn) and February (summer), but there was little reduction for lucerne. The peak leaf angle in the tedera accessions ranged from <40° to 70°, whereas for lucerne, the leaf angle was nearly parallel to incident light at 85°. Leaf water-use efficiency, relative leaf water content and leaf retention were higher for the tedera accessions than for lucerne in February. These results highlight the superior drought resistance of tedera compared with lucerne. The reduction in gs over the day in tedera shows the capacity of this species to reduce water loss quickly when conditions for CO2 fixation relative to water loss are highly unfavourable. The high retention of leaves in summer by tedera is a valuable trait for a perennial pasture plant in Mediterranean environments. Leaf folding, combined with effective stomatal control in summer, provides tedera with a set of physiological responses that confer high drought resistance.

Soilborne root disease pathogen complexes drive widespread decline of subterranean clover pastures across diverse climatic zones

Abstract. Subterranean clover (Trifolium subterraneum L.) is an important pasture legume in many regions of Australia, and elsewhere. A survey was undertaken in 2014 to define the levels of soilborne disease and associated pathogens in annual subterranean clover pastures across southern Australia. Most of the 202 samples processed had very severe levels of taproot rot disease (disease index 60–80%) and extremely severe lateral root rot disease (disease index 80–100%). A complex of soilborne root pathogens including Aphanomyces trifolii, Phytophthora clandestina, and one or more of Pythium, Rhizoctonia and Fusarium spp. was found responsible for severe pre- and post-emergence damping-off and root disease. This is the first study to highlight the high incidence of A. trifolii across southern Australian pastures and the first to highlight the existence of natural synergistic associations in the field between Rhizoctonia and Pythium spp., Pythium and Fusarium spp., Pythium spp. and A. trifolii, and P. clandestina and A. trifolii. Nodulation was generally poor, mainly only in the 20–40% nodulation index range. There was no relationship between rainfall zone and tap or lateral root disease level, with root disease equally severe in lower (330 mm) and higher (1000 mm) rainfall zones. This dispels the previous belief that severe root disease in subterranean clover is an issue only in higher rainfall zones. Although overall the relationship between tap and lateral root disease was relatively weak, these two root-disease components were strongly positively expressed within each pathogen’s presence grouping, providing explanation for variability in this relationship across different field situations where soilborne root disease is a major problem. Most producers underestimated the levels and effect of root disease in their pastures. This study established that tap and lateral root diseases are widespread and severe, having devastating impact on the feed gap during autumn–early winter across southern Australia. Severe root disease was independent of the highly variable complex of soilborne pathogens associated with diseased roots, geographic location and rainfall zone. It is evident that soilborne root diseases are the primary factor responsible for widespread decline in subterranean clover productivity of pastures across southern Australia. Implications for disease management and options for extension are discussed.