Latex is a sticky emulsion that exudes upon damage from specialized canals in about 10% of flowering plant species. Latex has no known primary metabolic function and has been strongly implicated in defense against herbivorous insects. Here we review historical hypotheses about the function of latex, evidence that it serves as a potent defense, and the chemistry and mode of action of the major constituent defense chemicals and proteins across a diversity of plant species. We further attempt to synthesize the characteristics of latex as a coordinated plant defense system. Herbivores that feed on latex-bearing plants typically evade contact with latex by severing the laticifers or feeding intercellularly, or may possess physiological adaptations. Convergent evolution appears to be rampant both in plants with latex and insects that exploit latex-bearing plants. Because latex shows phenotypic plasticity, heritability, and macoevolutionary lability, it is an ideal system to study plant-herbivore interactions using evolutionary approaches.

Since ancient times, mulberry leaves ( Morus spp.) have been used to rear the silkworm Bombyx mori . Because the silkworm grows well on mulberry leaves, the toxicities and defensive activities of these leaves against herbivorous insects have been overlooked. Here we show that mulberry leaves are highly toxic to caterpillars other than the silkworm B. mori , because of the ingredients of the latex, a milky sap exuded from mulberry leaf veins. The toxicity of mulberry leaves was lost when the latex was eliminated from the leaves, and artificial diets containing latex showed toxicity. Mulberry latex contained very high concentrations of alkaloidal sugar-mimic glycosidase inhibitors reported to have antidiabetic activities, such as 1,4-dideoxy-1,4-imino- d -arabinitol, 1-deoxynojirimycin, and 1,4-dideoxy-1,4-imino- d -ribitol. The overall concentrations of these inhibitors in latex reached 1.5-2.5% (8-18% dry weight) in several mulberry varieties, which were ≈100 times the concentrations previously reported from whole mulberry leaves. These sugar-mimic alkaloids were toxic to caterpillars but not to the silkworm B. mori , indicating that the silkworm can circumvent the mulberry tree's defense. Our results suggest that latex ingredients play key roles in defense of this tree and of other plants against insect herbivory, and they imply that plant latexes are treasuries of bioactive substances useful as medicines and pesticides.

Raphides, needle-shaped calcium oxalate crystals in tissues of many plants, have been thought to play defensive roles against herbivores without detailed bioassays for their defensive roles and modes of function using purified raphides. In order to examine the defensive roles and modes of function of raphides in a clear experimental system, we performed bioassays giving the larvae of the Eri silkmoth, Samia ricini (Saturniidae), leaves of their host plant, the castor oil plant, Ricinus communis (Euphorbiaceae), painted with the raphides purified from kiwifruits, Actinidia deliciosa (Actinidiaceae), in presence or absence of cysteine protease, which often coincide with raphides in plant tissues. Raphides alone or cysteine protease alone showed only weak defensive activities around experimental concentrations. However, when raphides and cysteine protease coexisted, they synergistically showed very strong growth-reducing activities, and the mortality of caterpillars was very high. In contrast, amorphous calcium oxalate did not show synergism with cysteine protease on defensive activities, indicating that the needle-shape of raphides is essential for the synergism. The present study provides the first clear experimental evidence for the synergism between raphides and other defensive factors. Further, the study suggests that “the needle effect”, which intensify the bioactivities of other bioactive factors by making holes to the barriers (cell membrane, cuticle, epithelium, the nuclear membrane, etc.) and facilitate the bioactive factors to go through them and reach the targets, is important in the defensive activities of raphides, and possibly in the allergy caused by raphides, and in the carcinogenic activities of other needle-shaped components including asbestos and plant derived silica needles.