Senescence-Associated Changes during Long-day-induced Leaf Senescence and the Nature of the Graft-transmissible Senescence Substance in Kleinia articulata

Schwabe, W. W. and Kulkarni, V. J. 1987. Senescence-associatedchanges during long-day-induced leaf senescence and the natureof the graft-transmissible senescence substance in Kleinia articulata.— J. exp. Bot. 38: 1741–1755. The long-day-induced senescence in Kleinia articulata leaveswas characterized by a loss in fresh and dry weight, in therate of leaf expansion and progressive loss of chlorophyll inthe detached rooted leaves. Ultrastructural examination of mesophyllcells of leaves from plants grown in continuous light showedthat osmiophilic globules accumulating in the chloroplasts werethe first visible sign of senescence in the organdies. Thesefirst signs of senescence could be detected in very young leavesof plants in continuous light, even before the leaves had expanded.Attempts were made to study the cause of this photoperiodicsenescence which, from previous work, appeared to involve agraft-transmissible substance. Leaves in continuous light showed reduced stomatal opening andextracts from them had very much higher activity in the Commelinastomatal closure assay (ABA-like activity ?) compared with non-senescingleaves grown in short days (8 h). However, even if all the activitywere due to ABA, this on its own does not appear to be the senescencesubstance because a much longer exposure to continuous lightwas required to induce irreversible senescence than to reachmaximum stomatal closure promoting activity in the bioassay.Moreover, severe water stress (high ABA?) did not lead to senescenceunless combined with continuous light or ethylene treatment.It is postulated that while ABA may play an important role inKleinia leaf senescence its lethal effect may not be realizedunless ethylene-induced membrane changes may synergisticallyassist. Key words: Leaf senescence, ABA, Daylength, stomatal movement, Kleinia     

Free Polyamines in Senescing Wheat(Triticum aestivum L.)

The free polyamine content of flag leaves, peduncles, rachis,glumes, and grains of wheat (Triticum aestivum L., cv. Castell)plants, ripening under field conditions, has been investigatedduring three consecutive growing seasons. Putrescine was quantitativelythe most important of all polyamines detected in these organs.Concentrations were highest in the grains, glumes and flag leaves.No correlation was found between polyamine content and the onsetof senescence of flag leaves and other organs. Excised primaryleaves, however, showed a decrease in polyamine content in thedark and also in light/dark cycles, but in the latter case onlyafter an initial increase. Sink removal of otherwise intactwheat plants caused an accumulation of putrescine in flag leavesat the later stages of senescence, whereas removal of all otherleaves was without any significant effect. Putrescine was alsorecovered in phloem-exudate samples collected throughout theperiod of grain development. In both grains and glumes, peakconcentrations of polyamines were found early during seed development. Key words: Triticum aestivum, polyamines, ripening, senescence     

NaCl-induced Senescence in Leaves of Rice (Oryza sativa L.) Cultivars Differing in Salinity Resistance

The influence of NaCl on senescence-related parameters (proteinand chlorophyll concentrations, membrane permeability and chlorophyllfluorescence) was investigated in young and old leaves of fiverice cultivars differing in salt resistance. NaCl hastened thenaturally-occurring senescence of rice leaves which normallyappears during leaf ontogeny: it decreased chlorophyll and proteinconcentrations and increased membrane permeability and malondialdehydesynthesis. Such an acceleration of deteriorative processes affectedall leaves in salt-sensitive cultivars while it was more markedin oldest than in youngest leaves of salt-resistant genotypes.NaCl-induced senescence also involved specific modifications,such as an increase in basal non-variable chlorophyll fluorescence(F ₀) recorded in all cultivars or a transient increase in solubleprotein concentration recorded in salt-resistant genotypes only.Alteration of membrane permeability appeared as one of the firstsymptoms of senescence in rice leaves and allowed discriminationamong cultivars after only 7 d of stress. In contrast, F _v/F _mratio (variable fluorescence/maximal fluorescence) was thesame for all cultivars during the first 18 d of stress and thuscould not be used for identifying salt-resistant rice exposedto normal light conditions. Relationships between parametersinvolved in leaf senescence are discussed in relation to salinityresistance of rice cultivars. Chlorophyll concentration; chlorophyll fluorescence; electrolyte leakage; magnesium; malondialdehyde; membrane permeability; NaCl; Oryza sativa L.; protein; rice; salinity resistance; senescence; UV absorbing substances     

A Vertical Gradient of the Chloroplast Abundance among Leaves of Chenopodium album

The abundances of chloroplasts in leaves on the main stems ofChenopodium album at different height levels were investigatedin relation to the photosynthetic capacity and light environmentof the leaves. (1) The number of chloroplasts per mesophyllcell decreased with descending position of leaves, except foryoung developing leaves at the top of plants that had smallerchloroplast numbers per cell than matured leaves beneath them.Contents of chlorophyll and ribulose-1,5-bisphosphate carboxylase/oxygenaseper leaf area that were highest in the topmost young leavesand decreased with decreasing height level indicate that thereis a vertical gradient of chloroplast abundance per leaf areadecreasing from the top of the leaf canopy with depth. (2) Light-saturatingrate of photosynthetic oxygen evolution per leaf area of maturedleaves decreased more steeply with decreasing leaf positionthan the chloroplast number per cell. Gradients of chlorophylland the enzyme protein contents were also steeper than thatof the chloroplast number. Loss of photosynthesis in lower leavesis, therefore, ascribed partly to loss of whole chloroplastsand partly to reduced photosynthetic capacities of the remainingchloroplasts. (3) The chloroplast number per cell in newly expandedsecond leaves was comparable to those in leaves that have developedat later stages of the plant growth but decreased graduallyduring leaf senescence both in the dark and light. The formationof the vertical gradient of chloroplast abundance is, therefore,ascribed to loss of whole chloroplasts during senescence ofleaves. (4) Irradiance a leaf receives decreased sharply fromthe top of the canopy with depth. The physiological or ecophysiologicalsignificance of the vertical distribution of chloroplasts amongleaves was discussed taking light environments of leaves intoconsideration. (Received July 31, 1995; Accepted October 20, 1995)     

Profiles of Leaf Senescence During Reproductive Growth of Sunflower and Maize

We investigated the effect of reproductive growth on the profilesof leaf senescence in maize (Zea mays L.) and sunflower (Helianthusannuus L.). Leaf senescence after flowering was assessed usingboth structural (leaf chlorophyll, nitrogen and dry matter)and functional (photosynthesis) variables in undisturbed plants(+G) and in plants in which grain set was prevented (-G). Twoweeks after flowering, lack of grain accelerated senescencein maize and delayed senescence in sunflower as indicated byleaf chlorophyll; leaf nitrogen and dry matter were less sensitiveresponse variables. Lack of interaction between reproductivetreatment and leaf position indicates that the senescence signal,whatever its nature, was equally effective throughout the plantin both species. In both species, feedback inhibition of photosynthesiswas first detected 30–35 d after flowering; excess carbohydratein the leaves was therefore an unlikely trigger of acceleratedsenescence in maize. As reproductive development progressed,differences between +G and -G plants were more marked in sunflower,and tended to disappear or reverse in maize. In sunflower, interactionsbetween leaf position and reproductive treatment—attributableto the local effect of grain—were detected around 20–27d after flowering. Copyright 2000 Annals of Botany Company Helianthus annuus, Zea mays, chlorophyll, light, nitrogen, photosynthesis, reproductive growth, senescence, source-sink, SPAD.     

The Relationship Between the Phosphorus Status of Subterranean Clover Plants and Losses in Chlorophyll from Detached Leaves: A Basis for the Visual Diagnosis of Phosphorus Deficiency

Leaves detached from subterranean clover plants (Trifolium subterraneumL.), grown in solution cultures at different phosphorus levels,were placed in water and in phosphate solutions. Losses in totalchlorophyll (a+b) of leaves in water were greater the lowerthe previous phosphorus supply to the plants from which theywere detached. In comparable leaves placed in phosphate solutionschlorophyll was maintained at levels at least as high as inleaves from non-deficient plants. For the latter there wereno differences between treated and untreated leaves. The differencesin colour change between treated and untreated leaves, therefore,increased with the deficiency in the phosphorus supply to theplants from which the leaves were detached, thus visibly reflectingtheir phosphorus status. A light intensity of 300 ft c (27 lx) or higher was necessaryfor maximum differences in leaf colour between treated and untreatedleaves, mainly because lower light levels reduced chlorophylllosses from leaves in water. Differences in colour between treated and untreated deficientleaves decreased with their age. However, valid comparisonsbetween treated and untreated leaf tissue could be ensured bythe use of treated and untreated leaflets. It is suggested that the results provide the basis for a simple,direct and visual diagnostic method, requiring no laboratoryequipment. Trifolium subterraneum L., subterranean clover, phosphorus deficiency, chlorophyll     

Influence of mycorrhizae and Rhizobium on cytokinin content in drought-stressed alfalfa

The objective of this research was to study the growth responseto drought of arbuscular mycorrhizal and non-mycorrhizal alfalfa(Medicago sativacv. Aragn) in relation to leaf cytokinin levels.In the experiment, four treatments were used: (a) plants inoculatedwith Clomus fasciculatum (Taxter sensu Gerd.) Gerdemann andTrappe and Rhizobium meliloti 102 F51 strain, (MR); (b) plantsinoculated with only Rhizobium (RP); (c) plants inoculated withonly mycorrhizae (MN); and (d) plants non-inoculated (NP). Non-mycorrhizalplants were supplemented with phosphorus and nonnodulated oneswith nitrogen to achieve similar size in all treatments. Plantswere subjected to drought by withholding irrigation in a cyclicway. The effects of drought on growth, number of stems, degreeof senescence, and leaf cytokinin levels were measured. Results of identification of cytokinins showed that dihydrozeatinriboside (dHZR) and ortno-topolin riboside (oTR) were predominantin alfalfa leaves. Nonsymbiotic plants (NP) showed higher totalcytokinin concentrations (dHZR and oTR). Under drought, NP plantsshowed the largest percentage drop in cytokinins and lower numberof stems as well as increased degree of senescent leaf tissuerelative to control values. By contrast, stressed symbioticplants (RP, MN and MR) showed higher green leaf weight thannonsymbiotic ones (NP) due to delay of leaf senescence and maintenance(RP) or increase (MN, MR) of stem leaf cytokinin levels duringdrought. The relationships between growth and the different cytokininsare discussed, suggesting an important role of mycorrhizal symbiosisin maintaining cytokinin levels under drought. Key words: Alfalfa, arbuscular mycorrhizae, cytokinins, drought, leaf senescence     

Water relations of cotton plants under nitrogen deficiency

Nitrogen deficiency in cotton plants (Gossypium hirsutum L.) increased the threshold water potentials for both stomatal closure and leaf senescence (defined as loss of chlorophyll and protein) during drought. These studies attempted to answer two questions: (1) What is the basis for the N/water interaction on senescence? (2) Is there a direct relationship between stomatal closure and senescence? Young and old leaves from N-deficient and N-sufficient plants maintained their relative sensitivities to water stress when excised leaf discs were floated on solutions of polyethylene glycol in dim light. In this leaf disc system, both leaf aging and N deficiency increased the threshold water potential for senescence. Leaf aging and N deficiency also decreased the concentration of exogenous abscisic acid necessary to initiate senescence in discs. A role for cytokinins in controlling senescence could not be clearly shown. In young leaves of both N-deficient and N-sufficient plants, stomata closed at water potentials much higher than those causing senescence. During leaf aging, the water potentials causing senescence increased more than those causing stomatal closure. The two processes thus occurred at about the same potentials in the oldest leaves. These data argue against a general cause-and-effect relationship between stomatal closure and senescence. Rather, each process apparently responded independently to absicsic acid accumulated during drought.     

The Effect of Age and Location of Leaf on Quantity and Quality of Japanese Mint Oil Production

Leaves of debranched Japanese mint (Mentha arvensis L., subsp.haplocalyx Briquet var. piperascens Holmes) were marked to allowsubsequent identification for age. Leaves of different age groupswere distilled for determination of quantity and quality ofoil. Most of the oil and its principal component, menthol, weresynthesized during the first 2 weeks of growth, while menthoneand the a-pinene percentage composition of the oil declinedwith age after 2 weeks. A significant decrease in percentageoil content occurred in older leaves indicating that leaf lossthrough senescence caused less loss of oil than of dry matter.Proportion of young to old leaves had a significant effect onoil composition. This is believed to be the first report onoil composition of leaves from single plants of Japanese mint.     

Apical Senescence in Pisum: A Direct or Indirect Role for the Flowering Genes ?

Apical senescence was examined in a range of intact and defloweredflowering genotypes under both long and short photoperiods.The flower inhibitor produced by the gene Sn, appears to havea direct effect on apical senescence since it can delay apicalsenescence under short day conditions in the absence of flowerand fruit development or where the rate of such developmentis the same in different treatments. Gene Hr can magnify thiseffect. Gene E, on the other hand, appears to influence apicalsenescence only indirectly through the effect it has on flowerand fruit development. The flowering genes at the If, sn andhr loci are also thought to have indirect effects on apicalsenescence. Even in deflowered plants apical senescence appearsto occur eventually in continuous light in all genotypes testedindicating that the presence of developing fruits, althoughpromotory, is not essential for apical senescence. Pisum sativum L., garden pea, flowering, senescence     

Can early wilting of old leaves account for much of the ABA accumulation in flooded pea plants?

When pea plants (Pisum sativum L.) were subjected to flooding,abscisic acid (ABA) content in shoots and roots increased upto 8-fold in the following days and stomatal conductance significantlydecreased. Although young leaves of flooded plants had a slightlyhigher water potential than those of the unflooded plants, oldleaves had lower water potential and lost turgor at the timewhen a substantial ABA increase was detected. In plants wherethe old leaves were clipped off, flooding did not cause anyABA increase during 7 d of the experimental period, except underconditions of higher transpiration demand, when the increasein ABA content was both delayed and small in scale (only I-fold).When intact plants were flooded and ABA was assayed separatelyin both old and young leaves, the ABA increase in old leavespreceded that in young leaves. Evidence here suggests that theflooding-induced ABA increase mainly results from the wiltingof old leaves. This suggests that young leaves may be protectedfrom wilting by ABA originating in old leaves under unfavourableenvironmental conditions. Key words: Waterlogging, soil flooding, ABA, leaf water relations, pea, Pisum sativum     

Leaf Water Diffusion Resistance in Clonal Tea (Camellia sinensis L.): Effects of Water Stress, Leaf Age and Clones

Leaf diffusion resistance was influenced by leaf age in well-wateredand water-stressed clonal tea plants. In well-watered plantsand in two of the three clones studied, young leaves showeda significantly lower diffusion resistance than old leaves.In water-stressed plants young leaves always exhibited a higherdiffusion resistance than old leaves. The highest diffusionresistance, irrespective of leaf age and water stress, occurredin clone DN which is known to be the most drought tolerant ofthe three clones studied, suggesting that drought tolerancein clone DN is caused, at least in part, by a stomatal regulationmechanism. Water release characteristic curves for the threeclones indicated differences in the water content-water potentialrelationship between young and old leaves as well as betweenclones. The drought tolerant clones had a higher relative watercontent for a given water potential compared with the drought-susceptibleclone. Camellia sinensis L., tea, diffussion resistance, water stress     

A Model of Leaf Tissue Growth, Acclimation and Senescence

A model for leaf growth, acclimation and senescence is described.The approach is mechanistic and the model, which is carbon-based,has eight state variables: substrate carbon and machinery carbon;primary, secondary and cross-linked cell wall; and two whichcorrespond to parameters of the photosynthesis light responsecurve—namely, the initial slope and light-saturated photosyntheticrate. The model is used to simulate responses to light and temperature,and predicts quantities such as final leaf area, the time atwhich the leaf becomes an exporter, senescence time, and thetwo photosynthetic parameters. The results are in good qualitativeagreement with observational responses that have been reported.The model also simulates the effects of changing the light environmentof a mature leaf: for instance, it predicts that increasingthe light level can increase leaf longevity and vice versa,although, in general, leaves grown continuously in high lighthave a shorter life than leaves grown continuously in low light. Leaf, model, growth, senescence, acclimation     

Nitrate Nutrition and Temperature Effects on Wheat: Enzyme Composition, Nitrate and Total Amino Acid Content of Leaves

Lawlor, D. W., Boyle, F. A., Kendall, A. C. and Keys, A. J.1987. Nitrate nutrition and temperature effects on wheat: Enzymecomposition, nitrate and total amino acid content of leaves.—J.exp. Bot. 38: 378–392. Wheat plants were grown in controlled environments in two temperatureregimes with two rates of nitrate fertilization. In some experimentstwo light intensities were combined with the nitrogen and temperaturetreatments. The composition of the third leaf was studied fromsoon after emergence until early senescence. The amounts ofchlorophyll, soluble protein, ribulose bisphosphate carboxylase-oxygenase(RuBPc-o) protein, nitrate, and total amino acids were measuredtogether with the activities of RuBPc-o, fructose- 1,6-bisphosphatase,glycolate oxidase, carbonic anhydrase, nitrate reductase, glutaminesynthetase and serine- and glutamate-glyoxylate aminotransferases.Additional nitrate supply increased the amounts, per unit leafarea, of chlorophyll, total soluble protein and RuBPc-o proteinand the activities of all the enzymes. The ratio of RuBP carboxylaseto RuBP oxygenase activity, when measured at constant CO²/O²ratio and temperature, was unaffected by growth conditions orleaf age. Leaves grown at the lower temperature, especiallywith more nitrate, contained much more soluble protein, nitratereductase, fructose bisphosphatase and free amino acids perunit area than the plants grown in the warmer conditions. However,young leaves grown in the warm contained more nitrate than thosegrown in the cool. Amounts of protein, amino acids and chlorophylland most enzyme activities reached maxima near full leaf expansionand decreased with age; additional nitrate slowed the decreaseand senescence was delayed. Nitrate content and nitrate reductaseactivities were highest in leaves before full expansion andthen fell rapidly after full expansion. Increased light intensityincreased the content of RuBPc-o protein at the higher rateof nitrate supply. Chloroplast components and, to a lesser extent,peroxisomal enzymes associated with photosynthetic nitrogenassimilation changed in proportion with different treatmentsbut nitrate reductase activity was not closely related to chloroplastenzymes. Control of tissue composition in relation to environmentalconditions is discussed. Key words: Nitrate nutrition, temperature, wheat, enzyme, amino acid, leaves, ribulose bisphosphate carboxylase oxygenase, nitrate reductase     

Selenium delays leaf senescence in oilseed rape plants

Effect of selenium on leaf senescence was studied in oilseed rape plants treated with 10 μM Na₂SeO₄ at a rosette growth stage. In addition to developmental senescence, N deficiency and leaf detachment were used for induction of senescence. Nonphotochemical quenching declined in old leaves as senescence became more advancing but rose progressively in the plants supplied by Se. The total carbohydrate and protein pools decreased with leaf age, while increased by the Se treatment. However, during senescence induced by N deficiency, Se did not change remarkably the C and N metabolism, but delayed senescence mainly through protection of plants from photoinhibitory effects. After detachment, untreated leaves became chlorotic and necrotic, while the Se-treated ones remained fairly green. Our results demonstrated that Se delayed leaf senescence by a maintaining or even improving photochemical activities. During developmental senescence, the Se effect on the extending life span of the leaves was additionally linked to the metabolic regulation of senescence.     

The Effects of Copper Supply and Shading on Retranslocation of Copper from Mature Wheat Leaves

Plants of Gamenya wheat (Triticum aestivum L.) were grown inpots of a Cu-deficient sand at two levels of Cu (deficient andsufficient), and harvested on days 13, 22, 28 and 38. In 50per cent of the pots in each Cu treatment, the oldest leaf andleaf 2 of the main stem were shaded when they reached full expansion. The Cu content of the oldest leaf of Cu-sufficient, unshadedplants was high at day 13 and declined rapidly to day 38. Thatof Cu-deficient, unshaded plants was initially relatively lowand declined much more slowly, so that at day 38 it resembledthat of Cu-sufficient plants. Shading the oldest leaf acceleratedthe loss of its Cu in both Cu-deficient and Cu-sufficient plants.The effects of shading and of Cu supply on the loss of Cu fromthe oldest leaf paralleled their effects on the loss of N andchlorophyll. The results suggest that most of the Cu in theoldest leaf does not move out until the leaf senesces. In Cu-deficient plants retention of Cu by old green leaves accentuatedCu deficiency. The release of Cu, resulting from shading theold leaves of Cu-deficient plants, stimulated the growth ofnew leaves. In Cu-sufficient plants, shading depressed growth. copper, shading, retranslocation, wheat, Triticum aestivum L.     

Effects of Abscisic Acid on Growth of Wheat (Triticum aestivum L)

Daily application of abscisic acid (ABA) to growing wheat plants,although initially inhibiting growth, resulted, after a shortlag, in an increase in the number of leaves and tillers. Thismay have been due to reduced apical dominance. At 84 days thetotal dry weight and area of all leaves produced up to thistime was less for the plants treated with ABA than for the controlplants. However, the area of green, living leaves and the dryweight were not significantly affected by the ABA treatment.Further effects of the daily ABA treatment were the inhibitionof transpiration, especially on the abaxial surface, the reductionof leaf size, the promotion of flowering and the stimulationof trichome formation on the leaf surfaces. ABA did not promoteleaf senescence in whole plants and actually increased leaflongevity. Triticum aestivum L., wheat, leaf senescence, transpiration, growth, flowering, abscisic acid     

Detection of Chlorophyll Breakdown Products in the Senescent Leaves of Higher Plants

The determination of intermediary breakdown products of chlorophyllsin senescent leaves of higher plants was performed by high-performanceliquid chromatographic technique that had been newly developedto separate and identify the oxidation products of chlorophylls,in particular monopyrrole derivatives [Suzuki et al. (1999)J. Chromatogr. A 839: 85]. In six plants tested, degradationproducts were detected only in the senescent leaves of plantsincluding barley (Hordeum vulgare), except for radish (Raphanussativus), in which they were found both in pre- and senescentcotyledons. In the senescent cotyledons of barley, three degradationproducts, hematinic acid, methyl ethyl maleimide, and the putativedegraded C-E-ring derivative, methyl vinyl maleimide dialdehyde,were detected. In addition to above three products, methyl vinylmaleimide was also found in both pre- and senescent cotyledonsof radish. These products decreased during senescence with anaccumulation of unknown compound(s), probably degraded monopyrrolederivatives. The degradation process and amounts of breakdownproducts of chlorophylls depend largely on plant species andvary with length of senescence. To conclude, it is likely thatchlorophylls are degraded into low-molecular-weight hydrolyticcompounds through monopyrroles. (Received March 10, 1999; Accepted June 11, 1999)     

The role of sugars in integrating environmental signals during the regulation of leaf senescence

Although leaf senescence results in a loss of photosynthetic carbon fixation, the senescence-dependent release of nutrients, especially of nitrogen, is important for the growth of young leaves and for reproduction. Environmental regulation of senescence is therefore a vital factor in the carbon and nitrogen economy of plants. Leaf senescence is a highly plastic trait that is affected by a range of different environmental factors including light, nutrient supply, CO2 concentration, and abiotic and biotic stress. In this review, the focus is on the impact of environmental conditions on sugar accumulation and sugar signalling during senescence. By signalling a high availability of carbon relative to nitrogen in the old leaves, sugar accumulation can trigger leaf senescence. Sugar-induced senescence is therefore particularly important under low nitrogen availability and may also play a role in light signalling. Whether or not sugars are involved in regulating the senescence response of plants to elevated CO2 remains unresolved. Senescence can be delayed or accelerated in elevated CO2 and no clear relationship between sugar accumulation and senescence has been found. Plasticity in the response to environmental factors, such as daylength and sugar accumulation, varies between different Arabidopsis accessions. This natural variation can be exploited to analyse the genetic basis of the regulation of senescence and the consequences for growth and fecundity. Different evolutionary strategies, i.e. early senescence combined with a high reproductive effort or late senescence combined with a low reproductive effort, may be an important adaptation of Arabidopsis accessions to their natural habitat.