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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.

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