Herbaspirillum-plant interactions: microscopical, histological and molecular aspects

Diazotrophic species in the genus Herbaspirillum (e.g. H. frisingense, H. rubrisubalbicans and H. seropedicae) associate with several economically important crops in the family Poaceae, such as maize (Zea mays), Miscanthus, rice (Oryza sativa), sorghum (Sorghum bicolor) and sugarcane (Saccharum sp.), and can increase their growth and productivity by a number of mechanisms, including nitrogen fixation. Hence, the improvement and use of these plant growth-promoting bacteria could provide economic and environmental benefits. We review the colonization processes of host plants by Herbaspirillum spp., including histological aspects and molecular mechanisms involved in these interactions, which may be epiphytic, endophytic, and even occasionally pathogenic. Herbaspirillum can recognize plant signals that modulate the expression of colonization traits and plant growth-promoting factors. Although a large proportion of herbaspirilla remain rhizospheric and epiphytic, plant-associated species in this genus are noted for their ability to colonize the plant internal tissues. Endophytic colonization by herbaspirilla begins with the attachment of the bacteria to root surfaces, followed by colonization at the emergence points of lateral roots and penetration through discontinuities of the epidermis; this appears to involve bacterial envelope structures, such as lipopolysaccharide (LPS), exopolysaccharide (EPS), adhesins and the type three secretion system (T3SS), but not necessarily the involvement of cell wall-degrading enzymes. Intercellular spaces are then rapidly occupied, proceeding to colonization of xylem and the aerial parts of the host plants. The response of the host plant includes both the recognition of the bacteria as non-pathogenic and the induction of systemic resistance to pathogens. Phytohormone signaling cascades are also activated, regulating the plant development. This complex molecular communication between some Herbaspirillum spp. and plant hosts can result in plant growth-promotion.