To cope with nutrient deficiencies, plants develop both morphological and physiological responses. The regulation of these responses is not totally understood, but some hormones and signaling substances have been implicated. It was suggested several years ago that ethylene participates in the regulation of responses to iron and phosphorous deficiency. More recently, its role has been extended to other deficiencies, such as potassium, sulfur, and others. The role of ethylene in so many deficiencies suggests that, to confer specificity to the different responses, it should act through different transduction pathways and/or in conjunction with other signals. In this update, the data supporting a role for ethylene in the regulation of responses to different nutrient deficiencies will be reviewed. In addition, the results suggesting the action of ethylene through different transduction pathways and its interaction with other hormones and signaling substances will be discussed.

Under Fe deficiency, Strategy I (non-graminaceous) plants up-regulate the expression of many Fe acquisition genes and develop morphological changes in their roots. The regulation of these responses...

• Asparagine metabolism in sunflower (Helianthus annuus) was investigated by cDNA cloning, sequence characterization and expression analysis of three genes encoding different isoforms of asparagine synthetase (AS, EC 6.3.5.4). • The AS-coding sequences were searched for in leaves, roots and cotyledons by using a methodology based on the simultaneous amplification of different cDNAs. Three distinct AS-coding genes, HAS1, HAS1.1 and HAS2, were identified. • HAS1 and HAS1.1 are twin genes with closely related sequences that share some regulatory features. By contrast, HAS2 is a singular sequence that encodes an incomplete AS polypeptide and shows an unusual regulation. The functionality of both the complete HAS1 and the truncated HAS2 proteins was demonstrated by complementation assays. Northern analysis revealed that HAS1, HAS1.1 and HAS2 were differentially regulated dependent on the organ, the physiological status, the developmental stage and the light conditions. • Asparagine synthetase from sunflower is encoded by a small gene family whose members have achieved a significant degree of specialization to cope with the major situations requiring asparagine synthesis.

This review deals with two essential plant mineral nutrients, iron (Fe) and phosphorus (P); the acquisition of both has important environmental and economic implications. Both elements are abundant in soils but are scarcely available to plants. To prevent deficiency, dicot plants develop physiological and morphological responses in their roots to specifically acquire Fe or P. Hormones and signalling substances, like ethylene, auxin and nitric oxide (NO), are involved in the activation of nutrient-deficiency responses. The existence of common inducers suggests that they must act in conjunction with nutrient-specific signals in order to develop nutrient-specific deficiency responses. There is evidence suggesting that P- or Fe-related phloem signals could interact with ethylene and NO to confer specificity to the responses to Fe- or P-deficiency, avoiding their induction when ethylene and NO increase due to other nutrient deficiency or stress. The mechanisms responsible for such interaction are not clearly determined, and thus, the regulatory networks that allow or prevent cross talk between P and Fe deficiency responses remain obscure. Here, fragmented information is drawn together to provide a clearer overview of the mechanisms and molecular players involved in the regulation of the responses to Fe or P deficiency and their interactions.

Iron (Fe) is an essential micronutrient for plants since it participates in essential processes such as photosynthesis, respiration and nitrogen assimilation. Fe is an abundant element in most soils, but its availability for plants is low, especially in calcareous soils. Fe deficiency causes Fe chlorosis, which can affect the productivity of the affected crops. Plants favor Fe acquisition by developing morphological and physiological responses in their roots. Ethylene (ET) and nitric oxide (NO) have been involved in the induction of Fe deficiency responses in dicot (Strategy I) plants, such as Arabidopsis. In this work, we have conducted a comparative study on the development of subapical root hairs, of the expression of the main Fe acquisition genes FRO2 and IRT1, and of the master transcription factor FIT, in two Arabidopsis thaliana ET insensitive mutants, ein2-1 and ein2-5, affected in EIN2, a critical component of the ET transduction pathway. The results obtained show that both mutants do not induce subapical root hairs either under Fe deficiency or upon treatments with the ET precursor 1-aminocyclopropane-1-carboxylate (ACC) and the NO donor S-nitrosoglutathione (GSNO). By contrast, both of them upregulate the Fe acquisition genes FRO2 and IRT1 (and FIT) under Fe deficiency. However, the upregulation was different when the mutants were exposed to ET [ACC and cobalt (Co), an ET synthesis inhibitor] and GSNO treatments. All these results clearly support the participation of ET and NO, through EIN2, in the regulation of subapical root hairs and Fe acquisition genes. The results will be discussed, taking into account the role of both ET and NO in the regulation of Fe deficiency responses.

Editorial: Nutrient Interactions in Plants