C. Christiansenweniger, ENDOPHYTIC ESTABLISHMENT OF DIAZOTROPHIC BACTERIA IN AUXIN-INDUCED TUMORS OF CEREAL CROPS, Critical reviews in plant sciences, 17(1), 1998, pp. 55-76
Gramineous crops such as wheat (triticum aestivum), maize (tea mays),
and rice (oryza sativa) develop tumorous structures (para-nodules) alo
ng primary and secondary roots when treated with low concentrations of
various auxins. Rice forms additional tumors along its hypocotyle. Hi
stologically, auxin-induced tumors appear as cancerous grown out root
meristems and thus are comparable in origin and structure to stem nodu
les of the legume sesbania rostrata. Auxin-affected root meristems do
not recover and:develop further to large nodule-like organs. Introduce
d diazotrophs (Azospirillum spp., Azorhizobium caulinodans, Rhizobium
spp.) potentially inhabit tissues of both stem and root tumors with th
e central meristem as a major colonization niche. Evidence is given th
at infecting bacteria follow a 'crack entry' invasion at sites where d
eveloping tumors have emerged through the root cortex and epidermis. B
acteria are shown to establish with high cell numbers inside intercell
ular spaces of cortical and meristematic tissues. Plant-cell infection
of tumor cells takes place with bacteria found inside the cell-cytopl
asm surrounded by membrane-like structures. Once inhabiting induced tu
mor tissues introduced diazotrophs colonize endophytically with high c
ell numbers. Mutant, ammonium-excreting and thus ecologically disadvan
taged A. brasilense is shown to survive inside para-nodulating maize a
nd rice plants with a dense population. Micro-aerobic nitrogenase acti
vities of tumor inhabiting diazotrophic bacteria (A. brasilense, Azoto
bacter vinelandii, A. caulonidans) are in general highly increased whe
n compared with untreated control plants. Additionally, bacterial nitr
ogenase activity is less sensitive to an increased oxygen tension in t
he root environment. The host plants benefit from the enhanced nitroge
n fixation in their para-nodulating roots. Highest rates of incorporat
ion of fixed nitrogen into host plant material is reported for para-no
dule inhabiting ammonium excreting A. brasilense strain C3. The host p
lant potentially stimulates the nitrogenase activity of endophytically
colonizing diazotrophs by providing energy in the form of a suitable
carbon source. In conclusion, it is demonstrated that gramineous plant
s are potentially capable of developing an endophytical diazotrophic s
ymbiosis through para-nodule formation.