Do silica-based defences drive plant-herbivore dynamics?

Understanding the factors that drive changes in the abundance of animal populations is fundamental to ecology. Many herbivore populations show regular oscillations in abundance, known as cycles, and these are usually thought to be due to matched oscillations in the abundance of predators rather than any changes in the herbivore's food plants. Food quality is not thought to respond to herbivory in a way which could lead to cycles, but we have discovered a novel way in which changes in plant quality could cause cycles in herbivore populations. This mechanism has not been considered before but it could apply to wide range of plant-herbivore systems. Our new idea is called the silica induction hypothesis. Periods of sustained heavy grazing lead to an increase in the levels of silica in grasses, so herbivores subsequently experience reduced availability of nutrients. This reduces their growth and reproductive rate and hence slows down the rate of population growth in the following year. Eventually populations fall to a level where there is only low grazing on the grasses, so the levels of silica in the leaves also fall because less well-defended leaves are produced. Herbivores are once again able to access nutrients in the grasses and their growth and reproduction increase again. We believe this mechanism can operate in many plant-herbivore systems, particularly ones based on grasses and other plants that contain high levels of silica. Silicon is the second most abundant mineral on earth and present in significant amounts in all plants, so the mechanism we propose is of wide relevance and significance. We already have some evidence from laboratory experiments and observations in the field that support our idea. In this project we aim to test this potential mechanism for the first time in large-scale field experiments. Firstly, we will determine the silica levels in grasses in areas where vole populations are high and compare them with those in areas where vole populations are low. If our ideas are correct, silica levels should be declining in areas where vole populations are increasing and vice versa. We will then set up an experiment to measure the rate and magnitude of the increase in silica at different levels of grazing and we will also measure how quickly the levels of silica defences decrease. Then we will test our ideas by moving voles into areas where we have induced high silica levels previously and see how feeding in these areas affects their growth and reproduction. These experiments will assess whether changes in plant defences can cause changes in herbivore abundance and help us develop a better understanding of the interactions between grasses and their herbivores. There are many important grassland systems that support a wide range of herbivores, including both rare species and livestock, so this project will be useful to both conservation and sustainable agriculture.