The Arabidopsis NRT1.1 transceptor coordinately controls auxin biosynthesis and transport to regulate root branching in response to nitrate

In agricultural systems, nitrate is the main source of nitrogen available for plants. Besides its role as a nutrient, nitrate has been shown to act as a signal molecule in plant growth, development, and stress responses. In Arabidopsis, the NRT1.1 nitrate transceptor represses lateral root (LR) development at low nitrate availability by promoting auxin basipetal transport out of the LR primordia (LRPs). Here we show that NRT1.1 acts as a negative regulator of the TAR2 auxin biosynthetic gene in the root stele. This is expected to repress local auxin biosynthesis and thus to reduce acropetal auxin supply to the LRPs. Moreover, NRT1.1 also negatively affects expression of the LAX3 auxin influx carrier, thus preventing the cell wall remodeling required for overlying tissue separation during LRP emergence. NRT1.1-mediated repression of both TAR2 and LAX3 is suppressed at high nitrate availability, resulting in nitrate induction of the TAR2 and LAX3 expression that is required for optimal stimulation of LR development by nitrate. Altogether, our results indicate that the NRT1.1 transceptor coordinately controls several crucial auxin-associated processes required for LRP development, and as a consequence that NRT1.1 plays a much more integrated role than previously expected in regulating the nitrate response of root system architecture.

The acropetal effects of indole-3-acetic acid in isolated shoot segments of Acer pseudoplatanus L. II. Possible regulation by a vectorial fieid of auxin waves

The acropetal effects of auxin on elongation of axillary buds and on modulation of the wave-like pattern of basipetal efflux of natural auxin to agar from <i>Acer pseudoplatanus</i> L. shoots were studied. When synthetic IAA was applied to cut surfaces of one of two branches the elongation growth of buds situated on the opposite branch was retarded, suggesting regulation independent of the direct action of the molecules of the applied IAA. Oscillations in basipetal transport of natural auxin along the stem segments were observed corroborating the results of other authors using different tree species. Apical application of synthetic IAA for 1 hour to the lateral branch caused a phase shift of the wave-like pattern of basipetal efflux of natural auxin, when the stem segment above the treated branch was sectioned. The same effect was observed evoked by the laterally growing branch which is interpreted as an effect of natural auxin produced by the actively growing shoot. These modulations could be propagated acropetally at a rate excluding direct action of auxin molecules at the sites of measurement. The results seem to corroborate the hypothesis suggesting that auxin is involved in acropetal regulation of shoot apex growth through its effect upon modulation of the vectorial field which arises when the auxin-waves translocate in cambium.

Auxin signal transduction into a quantitative change in natural auxin polar transport in pine cambium

Results of experiments performed with 6-mm high stem sections of <em>Pinus sylvestris</em> L. confirmed the hypothesis that the fusiform cells of the cambial region respond to the arrival of indole-3-acetic-acid (IAA) at the signalling concentration in the apoplastic space around their apical ends by increasing the basipetal efflux of endogenous natural auxin. Thus, the auxin signal propagation along the stem cambial region could be a chain of reactions between the axially neighbouring cells, each capable of responding and contributing to the change of auxin concentration in the apoplast which requires only transduction of the foreign auxin signal by each of the cells to increase the basipetal efflux of their own endogenous auxin. The rate of auxin signal propagation in this system is not limited by the rate of auxin molecular transport, and if it functions in conjunction with feedback inhibition, it may produce oscillations of the auxin basipetal efflux generating supracellular auxin waves. Inhibitors of the proteinaceous auxin efflux carrier associated with the plasmalemma (NPA and TIBA), although reducing total basipetal transport of the natural auxin, did not prevent the stimulated additional efflux of this phytohormone. The studied auxin-signal transduction processes seem to be intracellular but not mediated by the Ca-calmodulin complex. The natural auxin basipetal efflux increased significantly within 45 min following the period when it had been strongly reduced by successive collections to agar receivers replaced several times at the basal ends of 6-mm high stem sections in which the intact fusiform cells of the cambial region in about 90% are arranged in only one axial row. Such exhaustion of the cellular reserve of auxin did not prevent the additional auxin basipetal efflux stimulated by the IAA apical treatment. The results may suggest an effect of the auxin signal upon the supply of newly-synthesised auxin directly to the system responsible for its basipetal efflux.

LOP1: a gene involved in auxin transport and vascular patterning in Arabidopsis

We have taken a genetic approach to understanding the mechanisms that control vascular patterning in the leaves of higher plants. Here we present the identification and characterization of the lop1 mutant of Arabidopsis which is defective in basipetal transport of IAA. Mutant leaf midveins show disoriented axial growth, and bifurcation into twin veins that are frequently rotated out of the normal dorsal/ventral axis of the leaf. Mutant plants also display abnormal patterns of cell expansion in the midrib cortex and in the epidermis of the elongation zone of lateral roots. Lateral roots show abnormal curvature during initiation, sometimes encircling the primary root prior to growth in a normal downward direction. Mutant seedlings have normal levels of free IAA, and appear normal in auxin perception, suggesting that transport is the primary lesion. The abnormalities in vascular development, lateral root initiation and patterns of cell expansion observed in the lop] mutant are consistent with a basic disruption in basipetal transport of IAA.

752 PB 120 TRANSPORT OF TRYPTOPHAN AND CONVERSION TO IAA IN AVOCADO TISSUES

Tryptophan is known to be a precursor of IAA in plants. The amount of IAA available for the development of avocado fruit might be a limiting factor for its growth. It is well known that IAA is not transported into developing fruit along its strictly basipetal transport route. Therefore, IAA present in fruit must be synthesized in situ. We investigated the possibility that tryptophan or its metabolites are transported from leaf to fruit. An HPLC method was developed to quantitatively isolate and measure tryptophan and all well known intermediates in the synthesis of IAA. Avocado leaves were fed L-[side chain-3-14C] tryptophan and its transport and metabolism to IAA within the leaf and within the fruit were monitored over time. Significant movement of tryptophan or a metabolite from leaf to fruit occurs in 24 h.