Lateral organs are formed in plants by post embryonic developmental programs. Leaves, and flowers differentiate from the shoot apical meristem and lateral roots from the primary root pericycle meristem. Adventitious roots are roots formed from non-root lateral meristematic tissues, mostly the cambium, in many cases in response to stress signals. The ability of plants to regenerate adventitious roots is fundamental for selection and breading programs which are based on vegetative propagation of elite clones. Thus, recalcitrant plants, losing their rooting capability, may form a genuine commercial barrier in agricultural and forestry improvement programs. Some cellular mechanisms underlying adventitious root formation have been revealed, but much is yet to be clarified. The plant primary cell wall is a dynamic organ that can change its form, and perceive and relay molecular signals inward and outward during certain stages of development in particular cells. Therefore, before the secondary cell wall is deposited and plants become the wood from which walls and furniture are built, and the fibers from which cloths are woven, primary cell walls actively participate in plant cell differentiation and developmental programs. While auxin is a major regulator, cell walls are important in regulating coherent formative cell division and synchronized polar elongation of cell lineages that are necessary for lateral organ induction and formation, and collaborative cell functioning. Nevertheless, little is known of how cell wall changes are molecularly sensed and translated to intracellular signals during differentiation of adventitious roots. Here we summarize recent data linking, directly or indirectly, cell wall events to auxin signaling and to lateral or adventitious root induction and formation.

Summer and autumn in Israel are highly arid with not enough plants in bloom offering nectar and pollen to support the local apiary. This leads to decline in colony health and honey production. To increase food sources for honeybees, we initiated a project to clone elite Eucalyptus trees exhibiting constant and rich blooming from late summer to early winter. We induced adventitious roots from cuttings of two mature Eucalyptus trees of which nectar production and honeybees’ attraction was measured: Eucalyptus brachyphylla and Eucalyptus x trabutii. During the rooting process, a high frequency of cylindrical callus formation instead of roots was obtained. To shed light on the inner anatomy of the callus chunks, we compared their cell organization and cell-wall composition to those of roots. Whereas in the root, cells were organized in circumferential symmetry, no symmetry was found in the callus. Instead, a more chaotic accumulation of meristematic-like cells with sporadic clusters of tracheary elements laid in different directions were observed. The outer cell layer of the callus often included swollen cells with thin cell walls. Most callus cells stained more strongly for cellulose and lignin than cells in the root meristem. In addition, specific antibodies to methylesterified and de-methylesterified pectin showed differential staining of callus vs. root cells indicating cell wall differences. Strikingly, roots were seen to differentiate from the chaotic cell organization of the callus, albeit at low rates. Further investigation of the cellular and molecular mechanisms underlying callus formation, are required.

Dark-grown (etiolated) branches of many recalcitrant plant species root better than their green counterparts. Here it was hypothesized that changes in cell-wall properties and hormones occurring during etiolation contribute to rooting efficiency. Measurements of chlorophyll, carbohydrate and auxin contents, as well as tissue compression, histological analysis and gene-expression profiles were determined in etiolated and de-etiolated branches of the avocado rootstock VC801. Differences in chlorophyll content and tissue rigidity, and changes in xyloglucan and pectin in cambium and parenchyma cells were found. Interestingly, lignin and sugar contents were similar, suggesting that de-etiolated branches resemble the etiolated ones in this respect. Surprisingly, the branches that underwent short de-etiolation rooted better than the etiolated ones, and only a slight difference in IAA content between the two was observed. Gene-expression profiles revealed an increase in ethylene-responsive transcripts in the etiolated branches, which correlated with enrichment in xyloglucan hydrolases. In contrast, transcripts encoding pectin methylesterase and pectolyases were enriched in the de-etiolated branches. Taken together, it seems that the short de-etiolation period led to fine tuning of the conditions favoring adventitious root formation in terms of auxin–ethylene balance and cell-wall properties.