No abstract

Ammonium (NH4+) instead of nitrate (NO3-) as the nitrogen (N) source for tobacco (Nicotiana tabacum L.) cultivated in a pH-buffered nutrient solution resulted in decreased shoot and root biomass. Reduction of shoot fresh weight was mainly related to inhibition of leaf growth, which was already detectable after short-term NH4+ treatments of 24 h, and even at a moderate concentration level of 2 mM. Microscopic analysis of the epidermis of fully expanded leaves revealed a decrease in cell number (50%) and in cell size (30%) indicating that both cell division and cell elongation were affected by NH4+ application. Changes in various physiological parameters known to be associated with NH4(+)-induced growth depression were examined both in long-term and short-term experiments: the concentrations of total N, soluble sugars and starch as well as the osmotic potential, the apparent hydraulic conductivity and the rate of water uptake were not reduced by NH4+ treatments (duration 1-12 d), suggesting that leaf growth was neither limited by the availability of N and carbohydrates, nor by a lack of osmotica or water supply. Although the concentration of K+ in leaf press sap declined in expanding leaves by approximately 15% in response to NH4+ nutrition, limitation of mineral nutrients seems to be unlikely in view of the fast response of leaf growth at 24 h after the start of the NH4+ treatment. No inhibitory effects were observed when NH4+ and NO3- were applied simultaneously (each 1 mM) resulting in a NO3-/NH4+ net uptake ratio of 6:4. These findings suggest that the rapid inhibition of leaf growth was not primarily related to NH4+ toxicity, but to the lack of NO3(-)-supply. Growth inhibition of plants fed solely with NH4+ was associated with a 60% reduction of the zeatine + zeatine riboside (Z + ZR) cytokinin fraction in the xylem sap after 24 h. Furthermore Z + ZR levels declined to almost zero within the next 4 d after start of the NH4+ treatment. In contrast, the concentrations of the putative Z + ZR precursors isopentenyl-adenine and isopentenyl-adenosine (i-Ade + i-Ado) were not affected by NH4+ application. Since cytokinins are involved in the regulation of both cell division and cell elongation, it seems likely that the presence of NO3- is required to maintain biosynthesis and/or root to shoot transfer of cytokinins at a level that is sufficient to mediate normal leaf morphogenesis.

The present problems with hormonal signals transferring dominance effects are reviewed and, as a new hypothesis, it is stated that the sequence of sink development may establish the dominance effect. ‘Primigenic dominance’ (PD) is suggested to describe this kind of correlative inhibition, in which the earlier developed sink inhibits later developed organs. New results show that the polar IAA export of the earlier developed sink inhibits the IAA export of later developed sinks. This ‘autoinhibition’ occurs at ‘junctions’, where auxin streams from various sinks meet. It is suggested that this depressed IAA‐export of the subordinated fruit/sink acts as the signal that leads to inhibited development. This hypothesis avoids some of the problems related with other hypotheses, such as the requirement of a multidirectional signal. Primigenic dominance is a unifying hypothesis, which can be applied to most correlative dominance phenomena in the plant. Primigenic dominance is less complicated than the traditional ‘apical dominance’ (AD) hypothesis, because it does not require secondary messengers.

Leaf growth of many plant species shows rapid changes in response to alterations of the form and the level of N supply. In hydroponically-grown tomato (Lycopersicon esculentum L.), leaf growth was rapidly stimulated by NO(3)(-) application to NH(4)(+) precultured plants, while NH(4)(+) supply or complete N deprivation to NO(3)(-) precultured plants resulted in a rapid inhibition of leaf growth. Just 10 microM NO(3)(-) supply was sufficient to stimulate leaf growth to the same extent as 2 mM. Furthermore, continuous NO(3)(-) supply induced an oscillation of leaf growth rate with a 48 h interval. Since changes in NO(3)(-) levels in the xylem exudate and leaves did not correlate with NO(3)(-)-induced alterations of leaf growth rate, additional signals such as phytohormones may be involved. Levels of a known inhibitor of leaf growth, abscisic acid (ABA), did not consistently correspond to leaf growth rates in wild-type plants. Moreover, leaf growth of the ABA-deficient tomato mutant flacca was inhibited by NH(4)(+) without an increase in ABA concentration and was stimulated by NO(3)(-) despite its excessive ethylene production. These findings suggest that neither ABA nor ethylene are directly involved in the effects of N form on leaf growth. However, under all experimental conditions, stimulation of leaf growth by NO(3)(-) was consistently associated with increased concentration of the physiologically active forms of cytokinins, zeatin and zeatin riboside, in the xylem exudate. This indicates a major role for cytokinins as long-distance signals mediating the shoot response to NO(3)(-) perception in roots.

Response of cytokinin concentration in the xylem(Phaseolus vulgaris L.) plants to decapitation and and relationship to apical dominance exudate of bean auxin treatment, Abstract. When xylem exudate of previously untreated Phaseolus vulgaris plants was analysed for cytokinins by radioimmunoassay, a low concentration (about 5 ng-ml-L) was found. However, when the plants were decapitated about 16 h before the xylem exudate was collected, an almost 25-fold increase in cytokinin concentration was observed. Twenty-four hours after decapitation this increase even reached 4000% compared to control plants. Applying naphthaleneacetic acid (NAA) to the shoot of decapitated plants almost eliminated the effect of shoot tip removal on cytokinin concentration, suggesting that cytokinins in the xylem exudate of intact plants are under the control of the polar auxin transport system. Other xylem constituents, such as potassium or free amino acids did not show this strong increase after decapitation and did not respond to NAA application. It is concluded that the observed auxin/cytokinin interaction has an important regulatory role to play, not only in apical dominance but in many other correlative events as well.