Following cell division, fruit growth is characterized by both expansion through increases in cell volume and biomass accumulation in cells. Fruit growth is limited by carbon starvation; however, the mechanism controlling fruit growth under restricted carbohydrate supply is poorly understood. In a previous study using red-fleshed kiwifruit, we showed that long-term carbon starvation had detrimental effects on carbohydrate, anthocyanin metabolism, and fruit growth. To elucidate the mechanisms underlying the reduction in fruit growth during kiwifruit development, we integrated phytohormone profiling with transcriptomic and developmental datasets for fruit under high or low carbohydrate supplies. Phytohormone profiling of the outer pericarp tissue of kiwifruit showed a 6-fold reduction in total cytokinin concentrations in carbon-starved fruit, whilst other hormones were less affected. Principal component analysis visualised that cytokinin composition was distinct between fruit at 16 weeks after mid bloom, based on their carbohydrate supply status. Cytokinin biosynthetic genes (IPT, CYP735A) were significantly downregulated under carbon starvation, in agreement with the metabolite data. Several genes that code for expansins, proteins involved in cell wall loosening, were also downregulated under carbon starvation. In contrast to other fleshy fruits, our results suggest that cytokinins not only promote cell division, but also drive fruit cell expansion and growth in kiwifruit.

Fungal Epichloë endophytes form symbiotic associations with many temperate grasses, such as Lolium and Festuca, giving their host grasses an ecological advantage. The importance of specific Epichloë endophytes in providing varying levels of protection against invertebrate pests has been well documented. Similarly, but with fewer studies, the benefits of Epichloë to host grasses in drought events has been shown. Endophyte-infected grasses show an improved persistence against herbivore insect attack as well as resilience under drought. However, there are relatively few studies that investigate the interaction between drought and insect pressure, and yet it is these combined pressures that can prove detrimental for a ryegrass or fescue crop. This review examines the current state of knowledge on the effects of Epichloë on the interactions of insects and drought in temperate grasses.

Asexual Epichloë are obligate fungal mutualists that form symbiosis with many temperate grass species, providing several advantages to the host. These advantages include protection against vertebrate and invertebrate herbivores (i.e., grazing livestock and invertebrate pests, respectively), improved resistance to phytopathogens, increased adaptation to drought stress, nutrient deficiency, and heavy metal-containing soils. Selected Epichloë strains are utilised in agriculture mainly for their pest resistance traits, which are moderated via the production of Epichloë-derived secondary metabolites. For pastoral agriculture, the use of these endophyte infected grasses requires the balancing of protection against insect pests with reduced impacts on animal health and welfare.

Climate change is anticipated to lead to an increase in the occurrence and intensity of drought and fluctuations in insect cycles that will challenge modern pasture systems. Feeding by root aphids such as Aploneura lentisci Pass. can be a significant challenge to pastures. These below-ground living aphids are commonly found in New Zealand and Australia, feeding year-round on the roots of graminaceous plants such as perennial ryegrass (Lolium perenne L.). Some strains of the fungal endophyte Epichloë festucae var. lolii in perennial ryegrass can provide protection against root aphids and greater resilience under drought, contributing to higher persistency and growth than endophyte-free plants. However, the interaction between insect pressure and drought is not understood. This study examined the effect of drought on root aphid populations and plant performance in perennial ryegrass plants relative to endophyte status (±) and endophyte strain (AR37, NZ_CT) in a glasshouse experiment. Plants were cloned across the drought and well-watered treatments, and half of the plants were inoculated with root aphids, whilst half of the plants were treated with insecticide. Endophyte infection with strain NZ_CT and AR37 reduced root aphid numbers. Aphid populations were significantly higher in drought-stressed than in well-watered plants in both endophyte-infected and endophyte-free treatments. Under drought conditions, root aphid populations were increased 4-fold in NZ_CT and 8-fold in AR37 and endophyte-free plants in comparison with their well-watered counterparts. Root aphids reduced shoot dry weight by 16% in drought-exposed and 26% in well-watered plants in comparison with their insecticide-treated counterparts whilst reducing root biomass by 49%. Our results suggest that root aphids are likely able to exploit the higher availability of amino acids in the plant sap of drought-exposed plants. This study provides evidence that climate change-mediated impacts of root aphids could reduce production in perennial ryegrass-dominant pastures. However, field trials would be necessary to determine whether this effect is seen in situ, where numerous additional factors will be operating at the same time.