Fine structure of root hair infection leading to nodulation in the Frankia–Alnus symbiosis

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Abstract
Root hair infection by Frunkia (Actinomycetales) is the means by which nitrogen-fixing root nodules are initiated upon the actinorhizal host, Alnus rubra. Structural details of the infectious process and the changes in host root hair cells are demonstrated at the prenodule stage for the first time using light and transmission electron microscopy. The Frankia hypha is the infective agent, extending from the rhizosphere through the root hair wall in a highly deformed region of the hair. There is no evidence of pleomorphism of the Frankia hypha. The primary wall fibrils of the root hair appear disorganized at the site of penetration. There is extensive secondary wall formation in the infected hair. At the site of penetration, root hair cell wall ingrowths occur that are structurally consistent with transfer cell wall formation. The ingrowths are continuous with the encapsulating wall layer surrounding the Frankia hypha. The host cytoplasm is rich in ribosomes, secretory products, and organelles, including Golgi bodies, mitochondria, plastids, and profiles of endoplasmic reticulum. In an aborted infection sequence, some structural features of the host response to Frankia are observable, while other aspects of successful infection do not occur. Limited transfer cell wall is formed at the site of near infection. The root hair cytoplasm is senescent, however, and a callosic plug appears to surround the pathway of infection.