Everybody’s Dancing with Every One Else

A paper out of the Hawes Lab noted that root border cells respond to or probe the surrounding soil by releasing a multitude of proteins, amino acids, sugars and secondary metabolites, and they do so even after the cells detach from the root cap. This material, called exudates, appears to alter the surrounding rhizosphere community of microorganismsCurlango-Rivera et al., (2010) investigated the consequence of these plant exudates on the growth and production of root border cells. Exposure to pisatin, at concentrations capable of inhibiting fungal growth, stimulated production of root border cells, while exposure to a plant cell wall component, ferulic acid, inhibited growth of root border cells. Pisatin is a  phytoalexin, a secondary metabolite made by the plant to limit microbial attack.

Ferulic Acid
Ferulic Acid

Does this indicate that plant root investment in root border cells as sensory organs occurs under the protective influence of a plant phytoalexins, but that cell wall fragments resulting from the rupture of root cap cells under microbial attack may limit plant root border cell proliferation? I am curious about experimental results if simultaneous exposure to the compounds occurred? Ferulic acid is often bound to a matrix of polymer structures found in plant cell walls. Would it take time for diffusion of these compounds into the rhizosphere to gain concentrations able to impart a biological effect?

Plant Chemical Out Posts

Flower of Garden Strawberry (Fragaria ×ananassa)
Image via Wikipedia

The search for chemical mediators in plant root rhizosphere interactions with symbiotic and pathogenic organisms found in the soil continues to generate interesting research. Martha Hawes group at the University of Arizona reported on the role of sugars, proteins and small molecules found in root cap secretions – a mucilaginous mixture that covers the growing root tip and “converses” with the surrounding matrix of living organisms. The cap is rich in root border cells, which detach from the growing root tip. Curlango-Rivera et al (2010) provides us a bit more detail about which metabolites are biologically active. Neither sugars nor amino acids triggered root growth or border cell production. Transient exposure to biologically active concentration levels of the isoflavonoid pisatin, a phytoalexin, stimulated root border cell production but not root tip growth. I wonder if inhibition of root elongation may “reset” plant growth patterns as root border cells, acting as chemical sense organs, define the nature of the environment?

A second paper used histochemcial methods to profile root metabolites in plants from the Rose family (Hoffman et al., 2010). They found flavan-3-ol molecules in the root tip and border cells. Their findings suggest that the distribution of flavan-3-ols in Fragaria and Malus is under tight developmental control. These molecules are found in plants as catechin and epicatechin derivatives and in long chain (polymeric) form. They influence the taste and medicinal potential of green tea and wine, to name a few well-known plants. Previous researchers summarized their role in chelating toxic cations (metals) in the soil, establishing mycorrhizal interactions and priming plant root defense. This paper suggests a role in the transport of the long distance plant hormone auxin, which would link the chemical cross talk at root border cells with responses that occur in tissue distal to root tips. Hoffman’s research lacked a clear distinction of whether the monomeric or polymeric flavan-3-ol forms where the active species. This has plagued plant research for some time, since the analytical methods for detecting the polymeric forms have been crude and ineffective. All of their samples were from a botanical garden. I wonder if the flavan-3-ol profile would differ compared to native wild grown species?

References:

  1. Curlango-Rivera, G. et al. (2010) Plant Soil 332:267-275
  2. Hoffmann, T. et al. (2011) Plant Biology, 13: no. doi: 10.1111/j.1438-8677.2011.00462.x

Outpost Communication

Dr.  Martha Hawes has been a pioneering researcher on plant root border cells. I became fascinated with their role while researching the fungal/plant communication in the rhizosphere of goldenseal (Hydrastis canadendis) during my doctorate. I called her lab hoping someone might speak with me. She answered and spent an hour pointing out important research papers and suggesting approaches I might take to incorporate root border cell research. She was always open to helping anyone with a curious mind and passion for the subject into which she’d immersed her career efforts. I’m grateful to her for showing me generosity and kindness.

Plant root border cells are formed at the root tip where physical and biological interactions occur with the soil and microbe communities. The cells are genetically programmed to separate from the rest of the root structure and from each other. Cell-wall degrading enzymes dissolve cell wall matrix material that holds plant cells together. These “outpost” remain biologically active, excreting proteins and smaller molecules into the surrounding environment. Both types of molecules act as signals turning on/off gene expression to stimulate or prevent the growth of soil-borne bacteria and fungi. One important role appears to be in establishing a symbiotic relationship with mycorrhizal fungi (see previous post).

Few plants such as the Arabidopsis thaliana, which do not produce root border cells, also do not form mycorrhizal associations. In most plants, the content of border cells are accessible only to microorganisms able to recognize and respond to specific root signals. Among the compounds located in root border cells of various plants, medicinally valuable isoflavonoids modulate stable ecological relationships between mycorrhizal fungi and plant root tissue. These fungi stimulate the production of isoflavonoid in plant root tissue, while simultaneously the isoflavonoids increase mycorrhizal spore germination. The spores are an important survival mechanism used by the fungi. Measuring the activity in root border cells in “real time” as they interact with fungi is one of the great challenges to plant biologists.

Here’s a short video showing the release of border cells from a plant root cap:

More in-depth readings:
Harrison, M. and Dixon, R. (1993) Isoflavonoid accumulation and expression of defense gene transcripts during establishment of vesicular-arbuscular mycorrhizal associations in roots of Medicago truncatula. Mol. Plant Microbe Interact. 6:643-654
Hawes, M,C. et al (1998) Function of root border cells in plant health: Pioneers in the Rhizosphere, Annual Review of Phytopathology, 36:311-327.
Hawes, M.C. et al (2003) Root Caps and Rhizosphere. J. Plant Growth Reg. 21:353.
Kape, R. et al (1992) Legume root metabolites and VA-mycorrhiza development. J. Plant Physiol. 141:54-60.
Phillips D.A. et al (2004) Microbial products trigger amino acid exudation from plant roots. Plant Phys. 136: 2887-2894