Senescence in Leaves of Acer pseudoplatanus L. and Parthenocissus tricuspidata Planch: 1. Changes in Some Leaf Constituents during Maturity and Senescence

Data are recorded of the changes in chlorophylls, carotene,sugar, shikimic acid, and anthocyanin in leaves of sycamore(Acer pseudoplatanus L.) and Virginia creeper (Parthenocissustricuspidata Planch.) during leaf maturity and senescence. InParthenocissus the losses of chlorophylls, carotene, sugar,and shikimic acid during senescence were closely correlated,and were inversely related to the accumulation of anthocyanin.The losses of chlorophylls, carotene, and sugar by Acer leaveswere also closely correlated. No evidence was found to supportthe suggestion that anthocyanin formation was caused by accumulationof sugar during senescence, but marked differences in shikimicacid content were found between leaves of Acer, which did notform anthocyanin, and Parthenocissus. which did. It is suggestedthat autumn senescence of these leaves involves the rapid senescenceof an increasing proportion of the leaf tissue during a periodof 80 days, and that measurements of the content of constituentsgive an estimate of the proportion of the leaf tissue whichhas senesced.     

Graft Transmission of Longday-induced Leaf Senescence in Kleinia articulata

Kulkarni, V. J. and Schwabe, W. W. 1985 Graft transmission oflongday-induced leaf senescence in Kleinia articulata.—J.exp. Bot. 36: 1620–1633. Senescence of attached and detached rooted leaves in Kleiniaarticulata can be regulated by daylength, accelerated by longphotoperiods LD (> 16 h) and retarded by short days SD (8h). Using detached, rooted leaves as stocks in leaf to leafgrafts, senescence was readily transmitted from a LD donor leafto a SD receptor leaf even if the receptor was retained in SD(D8).However, no transfer could be detected where it had to passthrough any stem tissue. Senescence was reversible up to a certain stage, beyond whichgrafts senesced, pointing to an accumulation of the graft transmissiblefactor to a threshold level that causes irreversible death. Key words: Kleinia, leaf senescence, senescence factor, daylength, graft transmission     

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     

Senescence in Leaves of Acer pseudoplatanus L. and Parthenocissus tricuspidata Planch: II. Changes in Potassium and Sodium Content in Leaves and Leaf Discs of Acer

Changes in sodium and potassium content of the leaves of fiveAcer trees are recorded during two seasons. In general, sodiumcontents remained constant during leaf maturity and increasedduring senescence, whereas potassium contents increased duringleaf maturity and decreased during senescence. Short-term fluctuationsin sodium content during one season could be related to periodsof rainfall preceding some of the days of sampling. Experimentsin which excised leaves were washed in a spray of de-ionizedwater showed that the proportion of leaf potassium which couldbe removed increased as senescence progressed, and the quantityleached was approximately proportional to the area of the leafwhich had yellowed. With increasing leaf age, the amount of potassium which diffusedquickly out of discs cut from the leaves increased, whereasdiffusion of sodium became less. Discs cut from senescent leaveswere capable of absorbing sodium and it is suggested that thisability might account for the increase in sodium content ofAcer leaves during senescence.     

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     

Effect of Leaf Age and Shading on Photosynthesis in Potatoes (Solanum tuberosum)

The effects of leaf age and of shading on photosynthetic rateand on other leaf parameters of potato (Solanum tuberosum L.)were studied using a portable gas exchange system. A rapid decreasein the rate of photosynthesis during leaf senescence was observed.This was accompanied by an increase in stomatal resistance,and as a result a fairly constant level of sub-stomatal CO₂concentration was maintained at all leaf ages. The reductionin the photosynthetic rate in older leaves was therefore assumedto be essentially mesophyllic in origin, whereas the stomatalresponse was probably secondary. Canopy density significantly affected the rate of photosyntheticreduction with leaf age. Leaves maintained under high radiationintensities manifested a slower decline in their photosyntheticrate, especially in the early stages of their senescence, thanleaves kept under shade conditions. The latter leaves were foundto be more adapted to low radiation intensities, as indicatedby changes in their chlorophyll a:b ratio and specific leafweight Solanum tuberosum L, potato, photosynthetic rate, mesophyll, stomata, leaf age, radiation intensity, chlorophyll a:b ratio     

Intracellular Silica Deposition in Mature and Senescent Leaves of Sieglingia decumbens (L.) Bernh

Following 1, 2, 4, or 8 weeks of growth in a silica-minimalsolution, tillers of Sieglingia decumbens (Heath Grass) weretransferred for 8 days to a nutrient solution which contained50 or 100 ppm silica. The resultant formation of intracellularopal phytoliths (silica bodies) was compared for the four developmentalstages of leaf 3. Characteristically different phytolith typesoccurred in leaves of different ages. Senescent, in contrastto younger, mature leaves, typically exhibited extensive extracellularsilicification of the mesophyll, in addition to deposits instomatal, long, and bulliform cells of adaxial epidermis; depositionin abaxial silica cells and long cells, characteristic of youngleaves, was much reduced, or absent. Physico-chemical factors and cytoplasmic changes associatedwith senescence are discussed in relation to intracellular opalphytolith formation in mature leaves of S. decumbens. In thisrespect, the tendency of silicic acid sols to polymerize inthe presence of an organic matrix and mineral cations is consideredto be significant.     

Natural variation in the regulation of leaf senescence and relation to other traits in Arabidopsis

Leaf senescence results in the recycling of nutrients, thereby providing resources required for growth and reproduction. In this study, the effect of day-length on leaf senescence in eight different Arabidopsis thaliana ecotypes was determined and the relationship between senescence and other morphological and life history traits was analysed. A significant variation in the start and extent of leaf senescence depending on the genetic background and the response to day-length was found. Whereas senescence of early flowering ecotypes was accelerated by long days, no effect of day-length on senescence could be found in late flowering Kas-1 plants. Senescence in the different ecotypes was associated with other traits, such as floral transition, the total number of fruits, the total number of leaves and the maximum chlorophyll content. Plants that bolted early also senesced early, produced fewer leaves, accumulated less chlorophyll, but produced more fruits. The present results indicate that senescence may be a key component in the trade-off between investment in photosynthetic capacity and reproduction. The relationship between senescence and other traits was maintained independent of whether differences in senescence were caused by genetic (ecotype) or environmental (day-length) variation, suggesting that genetic and environmental factors affect these traits through common regulatory pathways.     

Chloroplast Protein Synthesis During Barley Leaf Growth and Senescence: Effect of Leaf Excision

Chloroplast protein synthesis was measured during the expansion,maturity and senescence of the oldest leaf of barley, Hordeumvulgare L., var. Hassan. A maximum rate of protein synthesisoccurred near the end of the expansion stage 9 d after sowing.Protein synthesis increased again at the beginning of senescenceand reached a new maximum at day 14 after sowing. Detachmentand incubation of leaves in the dark stimulated chioroplastprotein synthesis by fully expanded or by senescent leaves butnot by expanding leaves. If the detached leaves were kept inthe light, chloroplast protein synthesis was stimulated in fullyexpanded but not in senescent leaves. Short treatments (18 h)of leaf segments with growth substances in either light or indarkness, significantly changed the rate of protein synthesisshown by chloroplasts. The relationship between chloroplastprotein synthesis and leaf senescence is discussed. Key words: Hormones, light, maturity     

Mineral Depletion and Leaf Senescence in Soya Bean as Influenced by Foliar Nutrient Application During Seed Filling

Senescence, as judged by the time courses of leaf lamina photosynthesis,soluble protein and chlorophyll contents, was studied in relationto mineral redistribution in field-grown soya beans [Glycinemax (L.) Merr] to investigate the hypothesis that the depletionof nutrients m the leaves by the developing seeds is the causeof soya bean senescence. A mineral nutrient solution was appliedto the canopy during the seed-filling period, and the effectson senescence and mineral depletion of the leaves were determinedin three cultivars, at two leaf positions, weekly from beginningof seed filling through physiological maturity. The onset of senescence occurred shortly after the beginningof rapid seed filling Photosynthetic rate declined about 60per cent within 3 weeks. Protein dropped by 52 per cent andchlorophyll by 48 per cent over the same period. Foliar nutrient application, at a rate previously shown to givesignificant yield increases in soya beans, increased the concentrationsof N, P and K in the leaf laminae, but tended only to delaytheir decline and failed to either delay the onset or alterthe course of senescence. The results of this experiment seem to indicate that, undernormal growth conditions, the events of senescence in the soyabean are not causally related to the N, P or K concentrationsof the leaf laminae Glycme max (L.) Merr., soya bean, nitrogen, phosphorus, potassium, leaf protein, chlorophyll, photosynthesis, foliar nutrient application, mineral depletion, leaf senescence     

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     

Induction of leaf senescence by low nitrogen nutrition in sunflower (Helianthus annuus) plants

Different parameters which vary during the leaf development in sunflower plants grown with nitrate (2 or 20 mM) for a 42‐day period have been determined. The plants grown with 20 mM nitrate (N+) showed greater leaf area and specific leaf mass than the plants grown with 2 mM nitrate (N?). The total chlorophyll content decreased with leaf senescence, like the photosynthetic rate. This decline of photosynthetic activity was greater in plants grown with low nitrogen level (N?), showing more pronounced senescence symptoms than with high nitrogen (N+). In both treatments, soluble sugars increased with aging, while starch content decreased. A significant increase of hexose to sucrose ratio was observed at the beginning of senescence, and this raise was higher in N? plants than in N+ plants. These results show that sugar senescence regulation is dependent on nitrogen, supporting the hypothesis that leaf senescence is regulated by the C/N balance. In N+ and N? plants, ammonium and free amino acid concentrations were high in young leaves and decreased progressively in the senescent leaves. In both treatments, asparagine, and in a lower extent glutamine, increased after senescence start. The drop in the (Glu+Asp)/(Gln+Asn) ratio associated with the leaf development level suggests a greater nitrogen mobilization. Besides, the decline in this ratio occurred earlier and more rapidly in N? plants than in N+ plants, suggesting that the N? remobilization rate correlates with leaf senescence severity. In both N+ and N? plants, an important oxidative stress was generated in vivo during sunflower leaf senescence, as revealed by lipid peroxidation and hydrogen peroxide accumulation. In senescent leaves, the increase in hydrogen peroxide levels occurred in parallel with a decline in the activity of antioxidant enzymes. In N+ plants, the activities of catalase and ascorbate peroxidase (APX) increased to reach their highest values at 28 days, and later decreased during senescence, whereas in N? plants these activities started to decrease earlier, APX after 16 days and catalase after 22 days, suggesting that senescence is accelerated in N‐leaves. It is probable that systemic signals, such as a deficit in amino acids or other metabolites associated with the nitrogen metabolism produced in plants grown with low nitrogen, lead to an early senescence and a higher oxidation state of the cells of these plant leaves.     

Changes in Chlorophyll Content and Organization During Senescence of the Primary Leaves of Phaseolus vulgaris L. in Relation to Photosynthetic Electron Transport

A large decrease was observed in the chlorophyll content ofthe primary leaves of Phaseolus vulgaris during senescence.Chloroplasts isolated from mature and senescent leaves gavevery similar light saturation curves for electron transportreactions involving either PS I or PS II, indicating that theaverage number of chlorophyll molecules associated with eachreaction centre did not change during senescence. It is concludedthat the reaction centres ceased to function at the same timeas, or perhaps before, their antenna chlorophylls were lostfrom the thylakoid membrane, and that the percentage decreasein the number of functional reaction centres per leaf was atleast as great as the percentage decrease in the leaf chlorophyllcontent. The chlorophyll-protein composition of thylakoid membrane preparationswas examined by electrophoresis of samples treated with sodiumdodecyl sulphate. In older leaves a smaller proportion of thechlorophyll applied to polyacrylamide gels was associated withthe P700- chlorophyll a-protein complex. There was also a declinein emission at 734 nm in the 77 °K fluorescence spectrumof intact leaf tissue during senescence. These results indicatethat older leaves contained a smaller proportion of chlorophyllsassociated with PS I, and this is consistent with the decreaseobserved in the leaf chlorophyll a/b ratio during senescence.The effect of these changes in chlorophyll content on the capacityof the chloroplast to carry out photosynthetic electron transportis discussed.     

The Control of Flowering, Growth, and Dormancy in Kleinia articulata by Photoperiod

Kleinia articulata subjected to different photoperiods becomesdormant in long days (continuous light) and grows uninterruptedlyin short days (8 h). Dormancy results from the rapid cessationof leaf differentiation and expansion. By contrast stem growthis faster in long days and continues until all leaves have died.In long days growth is not halted permanently but flushes ofleaf production and stem elongation occur. Resumption of growthis stimulated rapidly by total darkness or short-day treatmentof the extreme stem tips for some days. This species is a strictshort-day plant in its flowering behaviour, depending on thisdaylength for normal inflorescence development also.     

Stomatal Responses and the Senescence of Leaves

Guard cell responses were examined in green and senescing leavesof Victa faba using detached epidermal strips to eliminate influencesfrom the mesophyll. Stomatal opening was greater in epidermalstrips from mature leaves than from senescing leaves althoughthe latter still retained the ability to respond to CO₂ andto kinetin. It was concluded that the decline in stomatal activityduring senescence is an independent but parallel process tochanges occurring in the mesophyll. Vicia faba, leaf senescence, stomata, kinetin     

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     

Effects of Leaf Age and Position on the Shoot Axis on Potassium Absorption by Tomato Leaf Slices

The effect of leaf age on K (⁸⁶Rb) influx into tomato (Lycopersiconesculentum Mill.) leaf lamina slices was determined for leaves5, 9 and 13 counting acropetally. Potassium influx rates expressedon a leaf fresh weight basis declined rapidly during leaf elongationat external KCI concentrations between 0.5 and 20.0 mM. In fullyexpanded leaves, K influx rates declined more slowly with age.The onset of senescence in mature leaves did not result in alarge loss in K uptake capability. Leaf position on the shootaxis and the stage of whole plant development had little influenceon K influx into leaf cells. It is suggested that the rapiddecrease in K influx in growing leaves is related to a dilutionin the concentration of K transporter sites resulting from anincrease in cell volume and weight. Lycopersicon esculentum Mill, tomato, free space, potassium, influx rate, ion uptake, leaf slices, leaf age leaf ontogeny     

Regulation of Leaf Senescence by Reproductive Sink Intensity in Cowpea (Vigna unguiculata L. Walp)

The relationship between seed number per pod and senescenceof the leaf in its axil was examined in a determinate cowpea(Vigna unguiculata L. Walp) variety C.779. The seed number perpod was reduced at all fruiting nodes by surgical excision ofpart of the 4-d-old pod. Leaf senescence as measured by lossof leaf area, chlorophyll content and soluble protein was sloweddown in leaves supporting the development of an artificiallyreduced number of seeds. Diminished nitrogen mobilization fromthe leaf could not account for the reduced rate of leaf senescence.The result suggests the involvement of a senescence signal fromthe developing seeds to the leaf in its axil. Development ofthe basal half of the excised pod in the cowpea provides a uniquesystem for manipulating seed number per pod. Senescence, monocarpic, chlorophyll, protein, Vigna unguiculata, cowpea     

Characterization of leaf senescence and pod development in soybean explants

Excised soybean (Glycine max [L.] Merrill) cv Anoka leaf discs tend to remain green even after the corresponding intact leaves have turned yello on fruiting plants. We have found that explants which include a leaf along with a stem segment (below the node) and one or more pods (maintained on distilled H₂O) show similar but accelerated leaf yellowing and abscission compared with intact plants. In podded explants excised at pre-podfill, the leaves begin to yellow after 16 days, whereas those excised at late podfill begin to yellow after only 6 days. Although stomatal resistances remain low during the first light period after excision, they subsequently increase to levels above those in leaves of intact plants. Explants taken at mid to late podfill with one or more pods per node behave like intact plants in that pod load does not affect the time lag to leaf yellowing. Explant leaf yellowing and abscission are delayed by removal of the pods or seeds or by incubation in complete mineral nutrient solution or in 4.6 micromolar zeatin. Like chorophyll breakdown, protein loss is accelerated in the explants, but minerals or especially zeatin can retard the loss. Pods on explants show rates and patterns of color change (green to yellow to brown) similar to those of pods on intact plants. These changes start earlier in explants on water than in intact plants, but they can be delayed by adding zeatin. Seed dry weight increased in explants, almost as much as in intact plants. Explants appear to be good analogs of the corresponding parts of the intact plant, and they should prove useful for analyzing pod development and mechanisms of foliar senescence. Moreover, our data suggest that the flux of minerals and cytokinin from the roots could influence foliar senescence in soybeans, but increased stomatal resistance does not seem to cause foliar senescence.