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P. Costantino - The role of the DAG transcription factors in Arabidopsis seed germination<br />
afb triple and quadruple mutants, anther dehiscence<br />
and pollen maturation occur earlier than in the wild<br />
type, causing the release of mature pollen grains<br />
before the completion of filament elongation. We<br />
also assessed the contribution of auxin transport to<br />
late stamen developmental processes. Our results<br />
suggest that auxin synthesized in anthers plays a<br />
major role in coordinating anther dehiscence and<br />
pollen maturation, while auxin transport contributes<br />
to the independent regulation of preanthesis filament<br />
elongation.<br />
Root meristem<br />
Plant postembryonic development takes place in<br />
the meristems, where stem cells self-renew and<br />
produce daughter cells that differentiate and give<br />
rise to different organ structures. For the maintenance<br />
of meristems, the rate of differentiation of<br />
daughter cells must equal the generation of new<br />
cells: how this is achieved is a central question in<br />
plant development. In the Arabidopsis root meristem,<br />
stem cells surround a small group of organizing<br />
cells, the quiescent center. Together they form<br />
a stem cell niche, whose position and activity<br />
depends on the combinatorial action of a small<br />
number of genes as well as on polar auxin transport.<br />
In contrast, the mechanisms controlling<br />
meristematic cell differentiation remain unclear.<br />
We demonstrated that cytokinins control the rate<br />
of meristematic cell differentiation and thus determine<br />
root meristem size via a two-component<br />
receptor histidine kinase-transcription factor signaling<br />
pathway. Analysis of the root meristems of<br />
cytokinin mutants, spatial cytokinin depletion, and<br />
exogenous cytokinin application indicated that<br />
cytokinins act in a restricted region of the root<br />
40<br />
meristem, where they antagonize a noncellautonomous<br />
cell-division signal, and we provided<br />
evidence that this signal is auxin.<br />
Subsequently, by means of a comprehensive genetic<br />
and molecular analysis, we showed that a primary<br />
cytokinin-response transcription factor, ARR1, activates<br />
the gene SHY2, a repressor of auxin signaling<br />
that negatively regulates the PIN auxin transport<br />
facilitator genes: thereby, cytokinin causes auxin<br />
redistribution, prompting cell differentiation.<br />
Conversely, auxin mediates degradation of the SHY2<br />
protein, sustaining PIN activities and cell division.<br />
Thus, the cell differentiation and division balance<br />
necessary for controlling root meristem size and root<br />
growth is the result of the interaction between<br />
cytokinin and auxin through a simple regulatory circuit<br />
converging on the SHY2 gene.<br />
Selected publications<br />
Dello Ioio R, Scaglia Linhares F, Scacchi E,<br />
Casamitjana-Martinez E, Heidstra R, Costantino P,<br />
Sabatini S. Cytokinins determine Arabidopsis root<br />
meristem size by controlling cell differentiation. Curr<br />
Biol. 2007, 17:678-82.<br />
Cecchetti V, Altamura MM, Falasca G, Costantino<br />
P, Cardarelli M. Auxin regulates Arabidopsis anther<br />
dehiscence, pollen maturation and filament elongation.<br />
Plant Cell 2008, 20:1760-74.<br />
Dello Ioio R, Nakamura K, Moubayidin L, Perilli S,<br />
Taniguchi M, Morita M, Aoyama T, Costantino P,<br />
Sabatini S. A genetic framework for the control of<br />
cell division and differentiation in the root meristem.<br />
Science 2008, 322:1380-4.