Pathological anatomy of the bacterial phloem canker disease of Juglans regia
Erwinia rubrifaciens Wilson, Zeitoun, and Fredrickson invades sieve tubes and parenchyma cells of the nonfunctional secondary phloem of Persian walnut, Juglans regia L. Because the sieve plate pores are great enough in diameter to allow passage of the bacteria, the nonfunctional phloem system provides an avenue along which the bacteria moves long distances up and down the bark. Functional phloem, on the other hand, does not exhibit symptoms of the disease nor is it found to contain the bacteria. Although the bacteria invade the ray parenchyma and move radially through these elements to the outer xylem, bacteria are not found to enter the xylem vessels. In culture, E. rubrifaciens produces long flexuous flagella. When taken from inoculated leaf tissue, however, it does not possess flagella. Hence, transport of bacteria from one sieve tube to another appears to be by apoplastic movement. Internal symptoms develop in two stages: a primary stage due to invasion of degenerate sieve tubes, and a secondary stage due to invasion of parenchyma cells and subsequent formation of wound callus. Pressure from wound callus induces vertical cracks in the bark. A slimy substance containing the bacteria exudes through the cracks to the bark surface, thereby allowing dispersion of the bacteria.