Plant growth and development involves a tight coordination of the spatial and temporal organization of cell division, cell expansion and cell differentiation. Orchestration of these events requires the exchange of signaling molecules between the root and shoot, which can be affected by both biotic and abiotic factors. The interactions that occur between plants and their associated microorganisms have long been of interest, as knowledge of these processes could lead to the development of novel agricultural applications. Plants produce a wide range of organic compounds including sugars, organic acids and vitamins, which can be used as nutrients or signals by microbial populations. On the other hand, microorganisms release phytohormones, small molecules or volatile compounds, which may act directly or indirectly to activate plant immunity or regulate plant growth and morphogenesis. In this review, we focus on recent developments in the identification of signals from free-living bacteria and fungi that interact with plants in a beneficial way. Evidence has accumulated indicating that classic plant signals such as auxins and cytokinins can be produced by microorganisms to efficiently colonize the root and modulate root system architecture. Other classes of signals, including N-acyl-L-homoserine lactones, which are used by bacteria for cell-to-cell communication, can be perceived by plants to modulate gene expression, metabolism and growth. Finally, we discuss the role played by volatile organic compounds released by certain plant growth-promoting rhizobacteria in plant immunity and developmental processes. The picture that emerges is one in which plants and microbes communicate themselves through transkingdom signaling systems involving classic and novel signals.

:  Melatonin (N‐acetyl‐5‐methoxytryptamine) is a tryptophan‐derived signal with important physiological roles in mammals. Although the presence of melatonin in plants may be universal, its endogenous function in plant tissues is unknown. On the basis of its structural similarity to indole‐3‐acetic acid, recent studies mainly focusing on root growth in several plant species have suggested a potential auxin‐like activity of melatonin. However, direct evidence about the mechanisms of action of this regulator is lacking. In this work, we used Arabidopsis thaliana seedlings as a model system to evaluate the effects of melatonin on plant growth and development. Melatonin modulated root system architecture by stimulating lateral and adventitious root formation but minimally affected primary root growth or root hair development. The auxin activity of melatonin in roots was investigated using the auxin‐responsive marker constructs DR5:uidA, BA3:uidA, and HS::AXR3NT‐GUS. Our results show that melatonin neither activates auxin‐inducible gene expression nor induces the degradation of HS::AXR3NT‐GUS, indicating that root developmental changes elicited by melatonin were independent of auxin signaling. Taken together, our results suggest that melatonin is beneficial to plants by increasing root branching and that root development processes elicited by this novel plant signal are likely independent of auxin responses.

N-acyl-homoserine lactones (AHLs) belong to a class of bacterial quorum-sensing signals important for bacterial cell-to-cell communication. We evaluated Arabidopsis thaliana growth responses to a variety of AHLs ranging from 4 to 14 carbons in length, focusing on alterations in post-embryonic root development as a way to determine the biological activity of these signals. The compounds affected primary root growth, lateral root formation and root hair development, and in particular, N-decanoyl-HL (C10-HL) was found to be the most active AHL in altering root system architecture. Developmental changes elicited by C10-HL were related to altered expression of cell division and differentiation marker lines pPRZ1:uidA, CycB1:uidA and pAtEXP7:uidA in Arabidopsis roots. Although the effects of C10-HL were similar to those produced by auxins in modulating root system architecture, the primary and lateral root response to this compound was found to be independent of auxin signalling. Furthermore, we show that mutant and overexpressor lines for an Arabidopsis fatty acid amide hydrolase gene (AtFAAH) sustained altered growth response to C10-HL. All together, our results suggest that AHLs alter root development in Arabidopsis and that plants posses the enzymatic machinery to metabolize these compounds.

Microorganisms and their hosts communicate with each other through an array of signals. The plant hormone auxin (indole-3-acetic acid; IAA) is central in many aspects of plant development. Cyclodipeptides and their derivative diketopiperazines (DKPs) constitute a large class of small molecules synthesized by microorganisms with diverse and noteworthy activities. Here, we present genetic, chemical, and plant-growth data showing that in Pseudomonas aeruginosa, the LasI quorum-sensing (QS) system controls the production of three DKPs-namely, cyclo(L-Pro-L-Val), cyclo(L-Pro-L-Phe), and cyclo(L-Pro-L-Tyr)-that are involved in plant growth promotion by this bacterium. Analysis of all three bacterial DKPs in Arabidopsis thaliana seedlings provided detailed information indicative of an auxin-like activity, based on their efficacy at modulating root architecture, activation of auxin-regulated gene expression, and response of auxin-signaling mutants tir1, tir1 afb2 afb3, arf7, arf19, and arf7arf19. The observation that QS-regulated bacterial production of DKPs modulates auxin signaling and plant growth promotion establishes an important function for DKPs mediating prokaryote/eukaryote transkingdom signaling.

Accumulating evidence indicates that plant growth promoting rhizobacteria (PGPR) influence plant growth and development by the production of phytohormones such as auxins, gibberellins, and cytokinins. Little is known on the genetic basis and signal transduction components that mediate the beneficial effects of PGPRs in plants. We recently reported the identification of a Bacillus megaterium strain that promoted growth of A. thaliana and P. vulgaris seedlings. In this addendum, the role of cytokinin signaling in mediating the plant responses to bacterial inoculation was investigated using A. thaliana mutants lacking one, two or three of the putative cytokinin receptors CRE1, AHK2 and AHK3, and RPN12 a gene involved in cytokinin signaling. We show that plant growth promotion by B. megaterium is reduced in AHK2-2 single and double mutant combinations and in RPN12. Furthermore, the triple cytokinin-receptor CRE1-12/AHK2-2/AHK3-3 knockout was insensitive to inoculation in terms of growth promotion and root developmental responses. Our results indicate that cytokinin receptors play a complimentary role in plant growth promotion by B. megaterium.