Plant secondary compounds and grasshoppers: Beyond plant defenses

Modern grasshoppers probably evolved from polyphagous ancestors endowed with the ability to tolerate many plant secondary compounds. This tolerance involves various behavioral and anatomical adaptations. Polyphagous grasshoppers have a relatively low level of sensitivity to the taste of many secondary compounds, and, if they do respond to the taste, have the capacity to habituate. This gives time for the induction of detoxifying enzymes so that unpalatable but potentially nutritious plants may be eaten safely. Associative learning involving secondary compounds may be important in food aversion learning, enabling the insects to avoid foods that have inappropriate nutrients, for example. Learning is also involved when grasshoppers develop associations between the taste of chemicals in the surface waxes of plants and internal leaf chemistry, enabling them to make Faster decisions about the acceptability of a plant. Anatomically, the midgut ceca of polyphagous grasshoppers have well-developed posterior arms, and it is possible that these are especially important in detoxification, while some species, in addition, have a specialized pocket region in which macromolecules accumulate to be eliminated from the body when the lining of peritrophic envelope is drawn out. Polyphagous species also have thick peritrophic envelopes to which various phenolics become adsorbed. Finally, the midgut environment contains surfactants that reduce tannin-protein complexing except at very high tannin concentrations. Some polyphagous species can utilize secondary compounds as defensive substances or, in one case, in cuticular sclerotization. Grass feeding has evolved on numerous occasions from these polyphagous ancestors, and it has: been associated with a loss of many of the characters providing protection from secondary compounds.