The involvement of abscisic acid (ABA) in the process of leaf abscission induced by 1 -aminocyclopropane-1 -carboxylic acid (ACC) transported from roots to shoots i n Cleopatra mandarin (Citrus reshni Hort. ex Tan.) seedlings grown under water stress was studied using norflurazon (NF). Water stress induced both ABA (24-fold) and ACC (16-fold) accumulation in roots and arrested xylem flow. Leaf bulk ABA also increased (&fold), although leaf abscission did not occur. Shortly after rehydration, root ABA and ACC returned to their prestress levels, whereas sharp and transitory increases of ACC (1 7-fold) and ethylene (1 O-fold) in leaves and high percentages of abscission (up to 47%) were observed. NF suppressed the ABA and ACC accumulation induced by water stress in roots and the sharp increases of ACC and ethylene observed after rewatering i n leaves. NF also reduced leaf abscission (7-1070). These results indicate that water stress induces root ABA accumulation and that this is required for the process of leaf abscission to occur. It was also shown that exogenous ABA increases ACC levels in roots but not in leaves. Collectively, the data suggest that ABA, the primary sensitive signal to water stress, modulates the levels of ethylene, which is the hormonal activator of leaf abscission. This assumption implies that root ACC levels are correlated with root
In this work, seedlings of two citrus rootstocks, the salt-tolerant Cleopatra mandarin (Citrus reshni Hort. ex Tan.) and the salt-sensitive Carrizo citrange (Citrus sinensis [L.] Osb. x Poncirus trifoliata [L.] Raf.) were used to study the relationship between chloride and water uptake. The results indicated that net chloride uptake rates in both genotypes were alike and decreased linearly with the time of salinity exposure, although they were more rapidly reduced in the tolerant genotype. In each rootstock, chloride uptake rates paralleled the decreases in transpiration rates. When transpiration was modified, concomitant changes in leaf Cl(-) concentrations were observed. There was a high positive correlation between total chloride content per plant and total water absorbed. In addition, the data indicate that the tolerant genotype "excluded" more chloride, i.e. it absorbed lower amounts of chloride per volume of water. Cleopatra also possessed a less efficient root system for water uptake and a higher shoot-to-root ratio. The results show that, overall, chloride absorption is linked to water use and that further tolerance in Cleopatra is mostly conferred by superior root resistance to Cl(-) uptake. Therefore, it is proposed that chloride absorption and, hence, salt tolerance in citrus depends to a great extent upon water use.
The hormonal signals controlling fruitlet abscission induced by sugar shortage in citrus were identified in Satsuma mandarin, Citrus unshiu (Mak.) Marc, cv. Clausellina and cv. Okitsu. Sugar supply, hormonal responses and fruitlet abscission were manipulated through full, partial or selective leaf removals at anthesis and thereafter. In developing fruitlets, defoliations reduced soluble sugars (up to 98%), but did not induce nitrogen and water deficiencies. Defoliation-induced abscission was preceded by rises (up to 20-fold) in the levels of abscisic acid (ABA) and 1-aminocyclopropane-1-carboxylic acid (ACC) in fruitlets. Applications to defoliated plants showed that ABA increased ACC levels (2-fold) and accelerated fruitlet abscission, whereas norflurazon and 2-aminoethoxyvinyl glycine reduced ACC (up to 65%) and fruitlet abscission (up to 40%). Only the full defoliation treatment reduced endogenous gibberellin A1 (4-fold), whereas exogenous gibberellins had no effect on abscission. The data indicate that fruitlet abscission induced by carbon shortage in citrus is regulated by ABA and ACC originating in the fruits, while gibberellins are apparently implicated in the maintenance of growth. In this system, ABA may act as a sensor of the intensity of the nutrient shortage that modulates the levels of ACC and ethylene, the activator of abscission. This proposal identifies ABA and ACC as components of the self-regulatory mechanism that adjusts fruit load to carbon supply, and offers a physiological basis for the photoassimilate competition-induced abscission occurring under natural conditions.
During ripening, citrus fruit-peel undergoes 'colour break', a process characterized by the conversion of chloroplast to chromoplast. The process involves the progressive loss of chlorophylls and the gain of carotenoids, changing peel colour from green to orange. In the present work, the in vivo and in vitro effects of supplemented nutrients (sucrose and nitrogen) and phytohormones (gibberellins [GA] and ethylene) on colour change in fruit epicarp of Satsuma mandarin (Citrus unshiu (Mak.) Marc., cv. Okitsu), were studied. The rate of colour break was correlated positively with sucrose content and negatively with nitrogen content. The removal of leaves blocked natural sucrose build-up and nitrogen reduction in the peel. Defoliation also inhibited chlorophyll disappearance and carotenoid accumulation, thereby preventing colour break. In vivo sucrose supplementation promoted sucrose accumulation and advanced colour break. In both in vivo and in vitro experiments, colour change promoted by sucrose was unaffected by ethylene but delayed by GA3. In non-supplemented plants, ethylene accelerated colour break while GA3 had no detectable effects. Ethylene inhibitors effectively counteracted the sucrose effects on colour change. Collectively, these results suggest that the chloroplast to chromoplast conversion in citrus fruit epicarps is stimulated by sucrose accumulation. The sugar regulation appears to operate via ethylene, whereas GA may act as a repressor of the sucrose-ethylene stimulation.
In citrus, the relative contributions of chloride and cations to growth disturbances induced by salinity are a matter of controversy. Chloride salts (15 mol m -3 CaCl 2 , 30 mol m -3 CaCl and 30 mol m -3 KCl) reduced growth and gas exchange parameters, increased leaf damage and abscission and produced anatomical disarrangements and mineral imbalances in seedlings of sensitive Carrizo citrange (Citrus sinensis x Poncirus trifoliata) and tolerant Cleopatra mandarin (Citrus reshni). In both cultivars, Ca 2+ was more beneficial, and K + more detrimental, for growth than sodium. Photosynthesis and growth disturbances were highly correlated (P ≤ 0·001) with leaf Clbuild-up. In the sensitive genotype, Cl -was also significantly correlated with several leaf anatomical disarrangements, such as increase in succulence. In comparison with sodium, both calcium and potassium increased leaf Clcontent (up to 25 and 69%, respectively). Protective calcium effects were not linked to improvement of photosynthesis, reduction of leaf anatomical disarrangements, or prevention of Cl -and Na + increases. It is proposed that the ameliorative effects of calcium on citrus grown under salinity are mostly related to reduction of leaf abscission. Collectively, the data suggest a cause-effect relationship between Cl -build-up and reduced growth, whereas chloride correlations with declines in photosynthesis or increases in succulence appear to be indirect.
The effect of water stress and subsequent rehydration on 1-aminocyclopropane-1-carboxylic acid (ACC) content, ACC synthase activity, ethylene production, and leaf abscission was studied in Cleopatra mandarin (Citrus reshni Hort. ex Tan.) seedlings. Leaf abscission occurred when drought-stressed plants were allowed to rehydrate, whereas no abscission was observed in plants under water stress conditions. In roots of water-stressed plants, a high ACC accumulation and an increase in ACC synthase activity were observed. Neither increase in ACC content nor significant ethylene production were detected in leaves of water-stressed plants. After rehydration, a sharp rise in ACC content and ethylene production was observed in leaves of water-stressed plants. Content of ACC in xylem fluid was 10-fold higher in plants rehydrated for 2 h after water stress than in nonstressed plants. Leaf (2,4,24). The relationship between ethylene production and abscission, both induced by water stress, has also been well established in cotton (11,15) and citrus (4). However, it has been indicated recently that these findings may be artifactual results obtained when working with detached tissues (19,20). These authors concluded that water stress alone cannot induce ethylene synthesis in intact plants.Citrus trees present a characteristic behavior of water stress-induced leaf abscission. When trees are subjected to water stress, leaves are injured but are not abscised. They remain attached until water stress is relieved (by rain or irrigation) and, soon afterwards, they are abscised (1). Similar behavior has been reported in cotton plants (15), which has led us to suggest that during water stress an abscissioninducing factor is synthesized in roots of citrus trees, and later it is transported to the aerial parts of the plant where it induces abscission. Ethylene is the natural regulator of abscission, and due to its gaseous nature, it may be that this abscission-inducing factor is ACC, the metabolic precursor of ethylene. Xylem transport of ACC from roots to shoots was first demonstrated by Bradford and Yang to induce epinasty in tomato plants subjected to anaerobic root stress (5, 6). Later, interorgan ACC transport was shown in several systems (3,9,12,18,21,23,28,29). However, no studies to test the relationship between water stress-induced abscission and xylem transport of ACC have been made. The aim of this work is to study this hypothesis. MATERIALS AND METHODS Plant MaterialEight-to 10-month-old Cleopatra mandarin (Citrus reshni TUDELA AND PRIMO-MILLO flux density, 120 Mmol-s-1 m2. RH was 95% during dark periods and oscillated between 60 and 95% during light periods. Plants were watered three times per week with 300 mL of half-strength modified Hoagland solution. Water Stress TreatmentsWater stress began at the start of the 16-h light period. Well-watered plants were taken from the pots and their roots washed with water. Then, the plants were placed in pots with dry sand for different periods under continuous light. In rehy...
Satsuma (Citrus unshiu [Mak] Marc.) and Clementine (Citrus reticulata [Hort.] Ex. Tanaka, cv Oroval) are two species of seedless mandarins differing in their tendency to develop parthenocarpic fruits. Satsuma is a male-sterile cultivar that shows a high degree of natural parthenocarpy and a high fruit set. Seedless Clementine varieties are self-incompatible, and in the absence of cross-pollination show a very low ability to set fruit. The gibberellins (GAs)
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