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Calcium, kinases and nodulation signalling in legumes

Key Points

  • Nod factors are oligosaccharide signalling molecules that are required for the establishment of nodule development. These signals activate several responses in the plant, including cytosolic Ca2+ fluxes and Ca2+ spiking in epidermal root-hair cells.

  • Mutations in a number of legume species define the genetic components that are involved in Nod-factor signalling. The proteins that are encoded by these genes can be placed in a signalling cascade relative to the cellular processes that are activated by Nod factor.

  • Some of the earliest-acting genes in this pathway encode receptor-like kinases with sugar-binding motifs in the extracellular domain. Mutations in these genes block all responses to Nod factor and this, coupled with the sequence homologies, suggests that their gene products probably function as the Nod-factor receptor.

  • Two genes downstream of the Nod-factor receptor are required for Ca2+ spiking and the maintenance of the Ca2+ flux. These genes encode a receptor-like kinase with leucine-rich-repeat domains in the extracellular domain and a protein with weak homology to cation channels.

  • Acting downstream of Ca2+ spiking, and potentially involved in the perception of this cellular response, is a chimeric Ca2+/calmodulin-dependent protein kinase. The recent cloning of the corresponding gene underlines the importance of Ca2+ in this signalling cascade.

Abstract

Several genes have recently been identified using legume mutants that are defective for nodulation signalling. The proteins they encode include novel types of receptor-like kinase that are predicted to recognize bacterial nodulation (Nod) factors, a leucine-rich-repeat receptor kinase, a putative ion channel and a predicted Ca2+/calmodulin-dependent protein kinase. The identification of these gene products provides new insights into the legume signalling responses to rhizobial signals.

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Figure 1: Legume infection by Rhizobium leguminosarum.
Figure 2: Structure of Nod factors.
Figure 3: Calcium spiking in a root-hair cell.
Figure 4: Predicted kinases that are required for Nod-factor signalling.
Figure 5: Orthologous legume genes and their predicted roles in early signalling events.
Figure 6: The Nod-factor signalling pathway in legumes.

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Acknowledgements

We would like to thank S. Walker for providing unpublished images and our colleagues for helpful discussions. The authors are supported by grants from the Biotechnology and Biosciences Research Council and the Royal Society (to G.E.D.O.).

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DATABASES

Interpro

EF hand

LysM

FURTHER INFORMATION

Lotus japonicus homepage

The World of Peas

Medicago truncatula Consortium

Glossary

BACTEROID

A terminally differentiated form of rhizobial bacteria that reside inside the nodule and fix nitrogen.

SYMBIOSOME

A bacteroid that is surrounded by a specialized plant membrane, and that is the site of nitrogen fixation and nutrient exchange.

FLAVANOID

A phenolic compound that is produced by the plant and that activates symbiotic responses in free-living rhizobial bacteria.

LysM DOMAIN

A domain that is proposed to be involved in binding β1–4-linked N-acetylglucosamine residues.

PEPTIDOGLYCAN

A proteinacious polysaccharide that is found in bacterial cell walls.

CHITIN

A polysaccharide that is made up of β1–4-linked N-acetyl glucosamine residues and is found in arthropod exoskeleton and some plants and fungi.

SYNTENIC

A region of the genome that is conserved between different species.

MYCORRHIZAE

Fungal species that form symbiotic interactions with plants and that assist in the uptake of nutrients from the soil.

BRASSINOSTEROIDS

A group of naturally occurring plant polyhydroxysteroids that function as plant hormones.

CLAVATA RECEPTOR

A receptor that is involved in meristematic identity.

VISININ

A Ca2+-binding protein in animals.

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Oldroyd, G., Downie, J. Calcium, kinases and nodulation signalling in legumes. Nat Rev Mol Cell Biol 5, 566–576 (2004). https://doi.org/10.1038/nrm1424

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