Possible Role Of Auxin Research Articles

Physiological Responses to Fe Deficiency in Split-Root Tomato Plants: Possible Roles of Auxin and Ethylene?

Iron (Fe) bioavailability in soils is often limited and can be further exacerbated by a non- homogeneous distribution in the soil profile, which has been demonstrated to vary both in space and time. Consequently, plants respond with morphological and physiological modifications at the root level involving a complex local and systemic signaling machinery. The present work unravels the role of two phytohormones (i.e., ethylene and auxin) and their integrated signaling in plant response to Fe deficiency. Inhibitors of auxin polar transport and of ethylene biosynthesis (N-1-naphthylphthalamic acid - NPA and aminoethoxyvinylglycine - AVG, respectively) were applied on tomato (Solanum lycopersicum L.) plants grown by the split-root technique, which allows to simulate condition of Fe heterogeneous distribution. Results showed that plants, exposed to an uneven Fe supply, triggered a complex auxin-ethylene signaling. A systemic action of auxin on FERRIC REDUCTASE OXIDASE 1 (SlFRO1) expression was revealed, while ethylene signaling was effective both locally and systemically. In addition, the investigation of Fe concentration in tissues showed that when leaves overcame Fe deficiency a Fe “steady state” was maintained. Therefore, physiological adaptation to this heterogeneous Fe supply could be mediated by the integration of the complex signaling pathways prompted by both auxin and ethylene activities.

Functional plasticity of miR165/166 in plant development revealed by small tandem target mimic

MicroRNA 165 and 166 (miR165/166) is composed of nine members and targets five members (PHB, PHV, REV, ATHB8 and ATHB15) of the HD-ZIP III transcription factor family. Mutants generated by traditional methods could hardly reveal the overall functions of miR165/166 in plant development. In this study, the expressions of all miR165/166 members were simultaneously blocked by over-expressing STTM165/166-31 in Arabidopsis and tomato for functional dissection of miR165/166 family. Following a down-regulation of over 90% endogenous miR165/166, the target HD-ZIP III genes were correspondingly up-regulated in the STTM transgenic Arabidopsis and tomato plants. Notably, the STTM165/166-31 over-expressed Arabidopsis and tomato displayed pleiotropic effects on development which were not frequently observed in previously identified genetic mutants of either individual miR165/166 gene or any of the five target genes. Furthermore, the transgenic Arabidopsis showed increased IAA content and decreased IAA sensitivity accompanied by enhanced expressions of genes responsible for auxin biosynthesis and signaling, suggesting possible roles of auxin in mediation of miR165/166-regulated processes. Importantly, the transgenic Arabidopsis exhibited the improved behavior under salt stress. Overall, such diverse variations in plant development and physiological process revealed by STTM165/166 demonstrate a key role of miR165/166-mediated network in regulating plant development and responses to abiotic stresses.

Auxin involvement in tepal senescence and abscission in Lilium: a tale of two lilies.

Petal wilting and/or abscission terminates the life of the flower. However, how wilting and abscission are coordinated is not fully understood. There is wide variation in the extent to which petals wilt before abscission, even between cultivars of the same species. For example, tepals of Lilium longiflorum wilt substantially, while those of the closely related Lilium longiflorum×Asiatic hybrid (L.A.) abscise turgid. Furthermore, close comparison of petal death in these two Lilium genotypes shows that there is a dramatic fall in fresh weight/dry weight accompanied by a sharp increase in ion leakage in late senescent L. longiflorum tepals, neither of which occur in Lilium L.A. Despite these differences, a putative abscission zone was identified in both lilies, but while the detachment force was reduced to zero in Lilium L.A., wilting of the fused tepals in L. longiflorum occurred before abscission was complete. Abscission is often negatively regulated by auxin, and the possible role of auxin in regulating tepal abscission relative to wilting was tested in the two lilies. There was a dramatic increase in auxin levels with senescence in L. longiflorum but not in Lilium L.A. Fifty auxin-related genes were expressed in early senescent L. longiflorum tepals including 12 ARF-related genes. In Arabidopsis, several ARF genes are involved in the regulation of abscission. Expression of a homologous transcript to Arabidopsis ARF7/19 was 8-fold higher during senescence in L. longiflorum compared with abscising Lilium L.A., suggesting a conserved role for auxin-regulated abscission in monocotyledonous ethylene-insensitive flowers.

In vitro testing of defense reactions in zygotic and somatic embryos of Abies numidica

Defense of desiccated cotyledonary somatic embryos and mature zygotic embryos of Abies numidica was tested in vitro by dual cultures with tester, fungus Phaeolus schweinitzii. Both types of embryos expressed defense reactions manifested by inhibited growth of fungal tester towards the embryos. Mycelial growth was described by logistic sigmoid growth model with a single asymptote. Mutual comparisons of mycelial growth in presence of zygotic and somatic embryos showed significant differences in parameters of mycelium growth curves towards the embryos. Larger defense reactions were observed in zygotic embryos relative to somatic embryos and unlimited control cultivations without embryo. The possible role of auxin in the defense response of plant embryos is discussed.

Possible roles of auxin and zeatin for initiating the dwarfing effect of M9 used as apple rootstock or interstock

Endogenous indole-3-acid (IAA) and zeatin (t-Z) may play important roles in the dwarfing mechanisms of rootstocks. The concentrations of IAA and t-Z, as well as the expression of the genes PIN1 and IPT3 were measured in leaves, barks, and roots from nine treatments: M9, Malus X micromalus Makino, Red Fuji/M9/Malus X micromalus, Red Fuji/M9, Red Fuji/Malus X micromalus, M9 rootstock substitution above and below the original graft union, interstock bridging, and interstock bark substitution of M9. The results show that there were greater amounts of t-Z and expressions of IPT3 in invigorated trees (Malus X micromalus and Red Fuji/Malus X micromalus) when compared with dwarfing trees (M9, Red Fuji/M9/Malus X micromalus and Red Fuji/M9) during the period of fast shoot growth (early June to mid-August). Moreover, the variation of IAA contents and PIN1 expressions shared the same pattern with t-Z contents and IPT3 expressions in all the tested locations. IAA content was extremely high in the bark of M9 interstock when compared with that of the scion and the rootstock, yet PIN1 expression in corresponding tissues was very low. After M9 rootstock were substituted above and below the original graft union, contents of t-Z, IAA, and PIN1 expressions in leaves and branch-barks recovered gradually to standard tree levels. However, there is no significant difference between the two treatments. We conclude graft union has no obvious influence on hormone transport. After M9 interstock and its bark were substituted, the hormone measurement of every index was consistent with that of rootstock substitution.

The interaction of light irradiance with auxin in regulating growth of Helianthus annuus shoots

Plants growing under canopy shade or in near-neighboring proximity of taller vegetation are the receivers of shade light conditions. The effect of light irradiance (photosynthetically active radiation [PAR]), one of the main components of shade light, on the growth of various tissues of sunflower seedlings and the possible role of auxin were investigated. Gradual reductions in PAR irradiance level from near-normal to low and very low result in significant and gradual increases in sunflower hypocotyl growth and endogenous auxin content. Similar reductions in PAR level resulted in significant and gradual decreases in sunflower cotyledon and leaf growth, and endogenous auxin content. Exogenously applied auxin increased hypocotyl elongation under near-normal PAR, where IAA levels are below optimum, but decreased elongation under very low PAR, where IAA levels are already at optimum. These results suggests that auxin acts as positive growth regulator of sunflower hypocotyls subjected to low light irradiance stress. This is further supported by the transfer experiments where seedlings transferred, for example, from near-normal PAR to very low PAR showed increased elongation associated with increased IAA levels. Therefore, it is reasonable to conclude that light irradiance-mediated changes in hypocotyl elongation of young sunflower seedlings are regulated by endogenous auxin levels.

Auxin and ethylene are involved in the responses of root system architecture to low boron supply in Arabidopsis seedlings

Changes in root architecture are one of the adaptive strategies used by plants to compensate for nutrient deficiencies in soils. In this work, the temporal responses of Arabidopsis (Arabidopsis thaliana) root system architecture to low boron (B) supply were investigated. Arabidopsis Col-0 seedlings were grown in 10 µM B for 5 days and then transferred to a low B medium (0.4 µM) or control medium (10 µM) for a 4-day period. Low B supply caused an inhibition of primary root (PR) growth without altering either the growth or number of lateral roots (LRs). In addition, low B supply induced root hair formation and elongation in positions close to the PR meristem not observed under control conditions. The possible role of auxin and ethylene in the alteration of root system architecture elicited by low B supply was also studied by using two Arabidopsis reporter lines (DR5:GUS and EBS:GUS) and two Arabidopsis mutants with impaired auxin and ethylene signaling (aux1-22 and ein2-1). Low B supply increased auxin reporter DR5:GUS activity in PR tip, suggesting that low B alters the pattern of auxin distribution in PR tip. Moreover, PR elongation in aux1-22 mutant was less sensitive to low B treatment than in wild-type plants, which suggests that auxin resistant 1 (AUX1) participates in the inhibition of PR elongation under low B supply. From all these results, a hypothetical model to explain the effect of low B treatment on PR growth is proposed. We also show that ethylene, via ethylene-insensitive 2 (EIN2) protein, is involved in the induction of root hair formation and elongation under low B treatment.

Nitrogen regulation of root branching.

Many plant species can modify their root architecture to enable them to forage for heterogeneously distributed nutrients in the soil. The foraging response normally involves increased proliferation of lateral roots within nutrient-rich soil patches, but much remains to be understood about the signalling mechanisms that enable roots to sense variations in the external concentrations of different mineral nutrients and to modify their patterns of growth and development accordingly. In this review we consider different aspects of the way in which the nitrogen supply can modify root branching, focusing on Arabidopsis thaliana. Our current understanding of the mechanism of nitrate stimulation of lateral root growth and the role of the ANR1 gene are summarized. In addition, evidence supporting the possible role of auxin in regulating the systemic inhibition of early lateral root development by high rates of nitrate supply is presented. Finally, we examine recent evidence that an amino acid, L-glutamate, can act as an external signal to elicit complex changes in root growth and development. It is clear that plants have evolved sophisticated pathways for sensing and responding to changes in different components of the external nitrogen supply as well as their own internal nitrogen status. We speculate on the possibility that the effects elicited by external L-glutamate represent a novel form of foraging response that could potentially enhance a plant's ability to compete with its neighbours and micro-organisms for localized sources of organic nitrogen.

Development of the adhesive pad on climbing fig (Ficus pumila) stems from clusters of adventitious roots

Vines have different climbing strategies; one type, called clinging vines, has developed a specialized structure, the adhesive pad that secretes a sticky substance that adheres to almost any substrate. In this study on the climbing fig, Ficus pumila L., we report on the developmental anatomy of clusters of adventitious roots that become transformed into this unique structure. Clustered adventitious roots in F. pumila are initiated in pairs on either side of a vascular bundle at the 2nd to 3rd internodes of young stems. After emergence through the cortex and epidermis, root hairs form, which secrete a substance that stains positively for polysaccharide and protein. The adventitious roots and root hairs stick together forming the adhesive pad. Pads stick to almost any substrate. If the adventitious roots do not touch a substrate they usually dry up, if they touch moist soil they tend to branch and change to a terrestrial form. We discuss experiments on the shoot developmental behavior leading to adhesive pad formation and the possible roles of auxin.

Plant Glutathione S-Transferases, a Tale of Theta and Tau

Glutathione S-transferases (GSTs, EC 2.5.1.18) are ubiquitous enzymes that catalyze the addition of glutathione to a wide variety of substrates. Plant GSTs have been studied mainly in relation to their role in the detoxification of herbicides, mostly in maize. Recently, several gst genes have been isolated as genes that respond to the plant hormone auxin with increased transcription. Pathogen infections and other treatments have also been found to lead to increased expression of gst genes. It is now apparent that different types of plant gst genes and activities exist in most plant species, but their functional relationships and their relationships to the action of auxin are still poorly understood. Here, a historic overview on plant GSTs will be presented. Based on primary sequence differences, a new class of plant GSTs, tau, will be proposed. The possible roles of auxin and oxidative stress in inducing gst genes will be discussed. Hopefully this review will help develop new ideas and stimulate new research to study the functions of the ever growing family of plant GSTs.

Characterization of two cDNAs that encode MAP kinase homologues in Arabidopsis thaliana and analysis of the possible role of auxin in activating such kinase activities in cultured cells.

Two cDNA clones, cATMPK1 and cATMPK2, encoding MAP kinases (mitogen-activated protein kinases) have been cloned from Arabidopsis thaliana and their nucleotide sequences have been determined. Putative proteins encoded by ATMPK1 and ATMPK2 genes, designated ATMPK1 and ATMPK2, contain 370 and 376 amino acid residues, respectively, and are 88.7% identical at the amino acid sequence level. ATMPK1 and ATMPK2 exhibit significant similarity to rat ERK2 (49%) and Xenopus MAP kinase (50%). The amino acid residues corresponding to the sites of phosphorylation (Thr-Glu-Tyr) that are involved in the activation of MAP kinases are conserved in ATMPK1 and ATMPK2. Northern blot analysis indicates that the ATMPK1 and ATMPK2 mRNAs are significantly present in all the organs except seeds. Genomic Southern blot analysis suggests that there are a few additional genes that are related to ATMPK1 and ATMPK2 in the Arabidopsis genome. Purified Xenopus MAP kinase kinase (MAPK kinase) phosphorylates ATMPK1 and ATMPK2 proteins that have been expressed in Escherichia coli, activating these enzymes. A rapid and transient activation of 46-kDa protein kinase activity that phosphorylated myelin basic protein (MBP) was detected when auxin-starved tobacco BY-2 cells were treated with synthetic auxin, 2,4-dichlorophenoxyacetic acid (2,4-D). Protein kinase activities which phosphorylated the recombinant ATMPK2 protein also increased rapidly after auxin treatment in the auxin-starved BY-2 cells. These results suggest that auxin may function as an activator of plant MAP kinase homologues, as do various mitogens in animal systems.

Stimulation of adventitious rooting in sunflower (Helianthus annuus) by low pH: possible role of auxin

Adventitious root formation by the hypocotyl cuttings of sunflower seedlings was greatly affected by the pH of buffered and unbuffered solutions bathing their basal portion. Exposure to low pH for 5 h after original root excision promoted root formation. Reduction of endogenous indole-3-acetic acid movement from the cotyledons and shoot apex was achieved by using N-1-naphthylphthalamic acid (an inhibitor of indole-3-acetic acid transport) and by removal of the cotyledons and shoot apex. Both the inhibitor and organ removal inhibited adventitious root formation, but acidic conditions could, to varying degrees, overcome this inhibition. Acidic conditions also increased the rate of [3H]indole-3-acetic acid uptake from the solutions around the hypocotyl bases and the rate of [3H]indole-3-acetic acid movement from cotyledons to the hypocotyl bases. Thus, acidic conditions may stimulate rooting by increasing the rate of basipetal indole-3-acetic acid transport to the zone of root initiation. These experiments show that in studies of the effects of various substances on rooting, the experimenter must be aware of these pH effects and take appropriate precautions. Key words: adventitious roots, auxin, indole-3-acetic acid, Helianthus annuus, pH.

The role of auxin in the removal of dormancy callose and resumption of phloem activity in Vitis vinifera

During winter the phloem of Vitis vinifera L. is dormant and is characterized by heavy deposits of callose on the sieve plates and lateral sieve areas of the sieve tubes. After bud break, the maturing leaves stimulate a nonpolar breakdown of this dormancy callose along the branch axis in the internodes located both above and below the leaves. However, the pattern of callose degradation in the radial direction is polar. It proceeds in a centrifugal direction so that the sieve tubes near the cambium become free of callose first and those adjacent to the periderm last. The effect of the leaves on the removal of dormancy callose can be replaced by auxin. Application of naphthaleneacetic acid to either the top or basal ends of excised dormant branches resulted in the removal of callose from sieve tubes, usually in less than a week. The fluorescent dye fluorescein was used to test phloem reactivation. Both acropetal and basipetal fluorescein movement occurred in sieve tubes in branches that were pretreated for 1 week with auxin, while much less movement of fluorescein occurred in the control branches, which remained dormant. Fluorescein translocation was observed in sieve tubes that had a reduced amount of callose and were wider than 20 μm, but was not detected in the narrow sieve tubes (diameters less than 15 μm) located next to the cambium. The possible roles of auxin, ethylene, and cytokinin in controlling callose levels in the sieve tubes are discussed. Key words: auxin, callose, fluorescein, Vitis vinifera, phloem, dormancy.

Senescence of Pear Fruit Cells Cultured in a Continuously Renewed, Auxin-deprived Medium

Auxins and cytokinins support cell division in tissue and cell cultures. In cytokinin-independent pear (Pyrus communis) cells, omission of 2,4-dichlorophenoxyacetic acid (2,4-D) from the medium for two successive transfers leads to rapid cell lysis, unless the osmolarity is raised to 0.4 molar with mannitol. Use of this system (nutrients plus mannitol minus 2,4-D) for the study of cell senescence was explored both in batch culture and in a system designed to permit medium renewal without withdrawal of live cells.In both systems, an initial period (1-6 days) of limited increase in cell number is characterized by a continuous decrease in the respiratory activity and in protein and RNA synthesis to very low basal rates. In batch culture, cell death occurs after 13 to 15 days with little or no change in metabolic activity, or in protein and RNA synthesis. With renewal of cell medium, death is slightly delayed and is preceded by a burst in RNA synthesis followed by a notable increase in protein synthesis. Cycloheximide inhibition of protein synthesis is transient and its effect on cell longevity variable. Nonetheless, in all instances cell death is preceded by a burst in protein synthesis. Actinomycin D (1.6 micromolar) did not significantly affect protein synthesis but delayed RNA synthesis and cell death. The possible roles of auxin, osmoticum, and macromolecular synthesis in cellular senescence and death are discussed.

Apical dominance in the tomato: The possible roles of auxin and abscisic acid

The roles of auxin (IAA) and abscisic acid (ABA) in controlling lateral bud development in young intact and decapitated tomato plants have been investigated using 2,3,5-triiodobenzoic acid (TIBA) as an inhibitor of the polar transport of apically-produced or exogenously-applied auxin. When TIBA was applied as a ring in lanolin to the stem, there was a strong stimulation of growth of the lateral buds below the ring, but these buds were suppressed in their outgrowth if either ABA was applied directly to them or the plants were grown under conditions which induce a high endogenous level of ABA. Such buds were suppressed whether or not the plants were treated with TIBA and only remained so as long as ABA was continuously supplied. It is suggested that the role of auxin may be one of inducing or maintaining a high level of ABA in the region of bud initiation and that it is this ABA which inhibits full bud development.

The Possible Role of Auxin in Reversing Photoperiodic Induction of Flowering in a Short-day Plant, Chenopodium rubrum L.

IAA (3.10 -4 M) inhibited the photoperiodic induction of flowering in the short day plant Chenopodium rubrum L. when applied during the inductive period to the shoot apex. When applied instead to the cotyledons it is much less effective, but only if the transport of auxin to the apex is inhibited by internode elongation due to gibberellic acid application. In some cases, as with TIBA treatment, this transport was reduced to less than 1 %. The rate of auxin transport was estimated using analytical methods or by semiconductography. The step-wise application of IAA during three or four inductive short days revealed that the inhibitory effect is restricted to the later stage of induction not inhibitable by actinomycine D or cycloheximide. The possible role of auxin in evocation is discussed.