Rhizobia colonize their legume hosts by different modes of entry while initiating symbiotic nitrogen fixation. Most legumes are invaded via growing root hairs by the root hair-curl mechanism, which involves epidermal cell responses. However, invasion of a number of tropical legumes happens through fissures at lateral root bases by cortical, intercellular crack entry. In the semiaquatic Sesbania rostrata, the bacteria entered via root hair curls under nonflooding conditions. Upon flooding, root hair growth was prevented, invasion on accessible root hairs was inhibited, and intercellular invasion was recruited. The plant hormone ethylene was involved in these processes. The occurrence of both invasion pathways on the same host plant enabled a comparison to be made of the structural requirements for the perception of nodulation factors, which were more stringent for the epidermal root hair invasion than for the cortical intercellular invasion at lateral root bases.ethylene ͉ host invasion ͉ nodulation factor ͉ symbiosis ͉ flooding-adapted growth L eguminous plants can engage in a symbiotic interaction with rhizobia (nitrogen-fixing bacterial symbionts), resulting in the formation of specialized root organs, the nodules. In the central nodule tissue, internalized bacteria fix dinitrogen to be used by the host. Many legumes are of agronomic importance as major food and feed crops, whereas others have a great potential as green manure. The legume-rhizobia interaction is initiated by a complex signal exchange during which recognition of bacterial nodulation (Nod) factors switches on the nodulation program in the plant. Nod factors are lipochitooligosaccharides that carry different substitutions. Recently, plant genes have been characterized that encode components of the Nod factor receptor͞ perception complexes (1-5).In legume symbiosis, bacterial invasion can follow different routes, the best known of which is via root hairs. Rhizobia induce the curling of growing root hairs, are entrapped in the curl, and enter the root hair by local hydrolysis of cell walls and invagination of the plasma membrane. Tip growth toward the base of the root hair results in an intracellular infection thread that proceeds through the cortical cells to reach the nodule primordia, where bacteria are released inside plant cells. The process takes place in the zone of developing root hairs (zone I, ref. 6), located just above the root meristem. Root hair invasion is used in pea, bean, soybean, vetch, and alfalfa, and in the model legumes Medicago truncatula and Lotus japonicus (7,8).Another mode of entry, via intercellular invasion at lateral root bases, has been observed in many tropical legumes. The bacteria enter via cracks formed by the protrusion of lateral roots and colonize large intercellular spaces called infection pockets. The mechanism for deeper invasion varies. In Sesbania rostrata and in Neptunia sp., infection pockets narrow down to form intercellular infection threads, and subsequently intracellular infection threads intrude in...