detect the presence of nematodes nearby in the soil and only builds its snares when they are present. This is presumably because building the net is a highly energy-consuming process; the fungus is alerted to the presence of the nematode by detecting the pheromones, such as ascarosides, with which the worms communicate. The fungus takes active steps to attract its prey by producing olfactory cues that mimic those used by the worm to find food and attract mates. Arthrobotrys dactyloides is a species that employs a loop of hypha to catch nematodes; when one tries to pass through the ring, the loop constricts with great rapidity, trapping the prey. Some nematophagous fungi produce toxic substances which immobilise nematodes. For example, the hypha of the shaggy ink cap (Coprinus comatus) attacks the free-living soil nematode Panagrellus redivivus with a structure known as a spiny ball; this is used to damage the nematode cuticle to enable immobilisation, after which the hypha pierces the skin and digests the contents. Most endoparasitic fungi have spores that are attracted to soil nematodes and tend to congregate in the mouth region. Having penetrated the cuticle, the hyphae grow throughout the nematode, absorbing its tissues. Escape tubes emerge from these and grow through the cuticle, and in due course, further motile spores exit through these, ready to infect other nematodes. In other species of fungi, it is conidia rather than spores which are encountered by the nematode and infect it in a similar way. In the case of Harposporium anguillulae, the sickle-shaped conidia are ingested by the nematode and lodge in the oesophagus or gut from where they invade the tissues. In egg parasitic species, the hypha flattens itself against the egg, the appearance of appressoria indicating that infection is about to take place or has already done so. The hypha then pierces the egg and devours the developing juvenile nematode before producing conidiophores and growing on towards nearby eggs. Biological control Some species of nematophagous fungi are being investigated for use in biological pest control. Purpureocillium lilacinum, for example, infests the plant-parasitic Meloidogyne incognita, which attacks the roots of many cultivated plants. Trials have provided varying results, with some strains being aggressive and others less pathogenic, and some strains that appeared promising in the lab proved ineffective in the field. Arthrobotrys dactyloides shows promise at controlling the cosmopolitan plant-parasitic root-knot nematode Meloidogyne javanica. References Bibliography Carnivorous